UNIVERSITY    OF    CALIFORNIA    PUBLICATIONS 

IN 

AGRICULTURAL    SCIENCES 

Vol.  1,  No.  5,  pp.  63-126  October  23,  1913 


STUDIES  UPON  INFLUENCES  AFFECTING 
THE  PROTEIN  CONTENT  OF  WHEAT 


BY 

G.  W.  SHAW 


In  1905  the  Merchants'  Exchange  of  San  Francisco,  the  State 
Board  of  Trade,  the  Sacramento  Valley  Development  Associ- 
ation, and  the  more  prominent  millers  and  grain  dealers  of 
California  called  the  attention  of  the  Agricultural  Department 
of  the  University  to  the  fact  that  the  milling  trade  found  it 
necessary  to  import  many  hundred  tons  of  wheat  per  year  to 
maintain  the  quality  of  California  flour  owing  to  the  low  gluten 
content  of  the  wheat  grown  in  the  state. 

With  the  view  of  ascertaining  the  causes  of  such  an  undesir- 
able condition,  investigations  were  begun  under  an  appropriation 
made  by  the  Legislature  at  the  session  of  1906  (Senate  Bill  no. 
10,  entitled  ''An  Act  to  provide  for  the  improvement  of  cereal 
crops  of  California  and  appropriating  money  therefor").  The 
investigations  are  still  under  way,  the  bill  having  been  re-enacted 
at  the  legislative  session  of  1908  and  again  in  1910. 

The  chief  points  for  study  in  these  investigations  are:  (1) 
To  determine  the  effect  of  changes  of  environment  upon  the 
growth  of  cereals,  particularly  as  regards  the  coinposition  of  the 
wheat  kernel  and  with  special  reference  to  the  causes  of  the 
production  of  a  low  protein  content;  (2)  to  discover  o»  produce 
such  wheat  as  will  yield  the  largest  profit  per  acre  for  the  farmer, 
and  will  supply  the  millers  with  wheat  of  superior  quality; 
(3)  to  conduct  similar  experiments  with  oats,  barley,  and  other 
cereals  as  may  he  desirable;  (4)  to  determine  the  effectiveness  of 
various  methods  of  culture  as  affecting  the  production  of  cereals. 


64  University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 

The  nature  of  this  work  is  such  that  no  permanent  results 
can  be  secured  from  one  or  two  seasons'  operations.  The  element 
of  time  is  an  all-essential  one  for  the  solution  of  such  problems. 
This  would  be  true  even  though  the  problem  was  merely  the 
development  of  wheats  giving  higher  yields  than  those  now  being 
gro^^^l  in  the  State,  but  ivith  the  introduction  of  the  problem  of 
increasing  the  ghden  content,  the  element  of  time  is  even  of 
greater  importance.  With  this  in  view,  the  experiments  were 
so  planned  that  while  the  final  results  could  not  be  expected  for 
several  years,  it  was  possible  to  obtain  data  of  importance  bearing 
upon  the  entire  question  of  cereal  culture  each  year,  material 
progress  thus  being  assured. 

As  early  as  1882  Clifford  Richardson  called  attention  to  the 
fact  that  wheats  from  the  Pacific  Coast  were  relatively  low  in 
their  protein  content,  and  numerous  analyses  of  California 
grown  wheats  made  at  the  California  Agricultural  Experiment 
Station  under  the  direction  of  Dr.  E.  W.  Hilgard,  in  the  earlier 
years,  also  showed  the  same  condition  to  exist. 

That  this  condition  exists  is  undoubtedly  true,  whatever  the 
causes  may  be.  To  bear  witness  to  the  fact  the  following  sum- 
mary is  given  of  analyses  of  white  wheats  grown  in  several 
different  states,  as  compared  with  the  anal3^ses  of  149  samples 
of  wheats  of  the  same  class  grown  in  California  in  the  same 
years.  For  the  sake  of  comparison  also  the  average  results 
obtained  from  the  analysis  of  49  hard  winter  wheats  grown  in 
Kansas  the  same  season  are  included,  as  well  as  those  of  three 
soft  white  wheats  grown  in  Kansas  from  seed  originally  obtained 
from  the  state  of  Washington. 

Table  Showing  the  Protein  Content  of  White  Wheats  Grown  in 
Different  States  in  the  Same  Season 

Number  Total 


State 

of 
samples 

protein 
Dri/  basis 

California 

149 

9.77 

Idaho 

46 

10.88 

Nevada 

34 

16.18 

Montana 

6 

14.24 

Utah 

10 

15.20 

Washington 

177 

1 2.98 

Kansas  (liard 

winter) 

49 

13.2.5 

Kansas  (white 

wheats)  * 

3 

10.57 

Grown   in   Kansas  from  seed  secured  from  the  State  of  Washington. 


1913]     Shaiv :  Studies  on  Infmences  Afecting  Protein  Content  of  Wheat        65 

In  practically  all  of  the  former  studies  of  this  subject  the 
plan  has  been  to  transfer  the  seed  from  one  point  to  another  and 
thus  change  the  environmental  conditions,  and  from  the  results 
so  secured  to  attempt  to  draw  general  conclusions.  Under  such 
conditions,  changes,  not  merely  in  environment,  but  also  in  soil, 
were  accomplished,  thus  introducing  too  many  variables.  In 
the  experiments  here  recorded  two  general  methods  have  been 
followed:  {a)  the  production  of  numerous  varieties  of  wheat  in 
the  field  on  a  small  area  of  uniform  soil  and  varying  other  con- 
dition of  growth  than  the  soil,  and  (5)  growing  wheat  from  the 
same  seed  under  the  same  conditions  on  soils  of  widely  different 
origin  placed  under  the  same  cultural  and  climatic  influences. 
The  large  and  important  question  involved  is  the  cause  or  causes 
of  the  relatively  low  gluten  content  of  wheat  grown  under  Cali- 
fornia conditions.  In  this  connection,  it  is  particularly  desirable 
to  know  whether  or  not  such  tendency  to  change  as  exists  is 
constant;  whether  it  is  due  to  some  climatic  influence,  the  time 
of  cutting,  the  time  of  seeding,  the  bleaching  action  of  the  sun, 
the  efl^ect  of  early  and  late  application  of  moisture,  the  effect 
of  cold  nights,  of  varying  amounts  of  sunshine  during  the  rip- 
ening period,  or  to  some  induced  or  inherent  condition  of  the 
soil.  The  work  which  we  have  conducted  has  aimed  at  the  solu- 
tion of  all  of  the  questions  just  mentioned  and  consisted  of 
several  parts  each  of  which  dealt  with  one  of,  the  questions 
involved. 

Variation  in  the  Protein  Content  of  Wheat 

Seasonal  Variation. — It  has  been  observed  that  from  season 
to  season  there  is  a  marked  variation  in  the  protein  content  of 
wheat  even  though  grown  upon  the  same  soil,  a  seeming  indi- 
cation that  the  seasonal  factor  is  considerably  greater  than  the 
soil  factor  in  protein  formation.  This  may  be  shown  from  a 
number  of  samples  grown  during  the  course  of  the  experiments, 
some  of  which  are  given  below.  These  wheats  have  been  grown 
each  year  on  the  same  soil  and  under  the  same  cultural  conditions 
at  the  University  Farm  at  Davis,  California. 


University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 


Total  Protein  in  Dry  Matter 

1906      1907     1908      1909      1910  1911  1912 

Kubanka  9.68  9.90         14.71         14.30         14.87  10.60  14.93 

Crimean  10.38         11.38         11.75         13.27         12.18  10.60  12.11 

Little  Club  Av.  of  5  plats  10.29  9.78  9.98  13.00 


That  the  quantity  of  available  nitrogen  in  the  soil  has  com- 
paratively little  bearing,  if  any,  beyond  its  necessity  in  sufficient 
quantity  to  insure  the  normal  growth  of  the  plants,  is  indicated 
on  a  series  of  fertilizer  plats  of  very  uniform  land,  discussed 
further  on  in  this  paper.  The  seasonal  effect  is  apparent,  how- 
ever, from  the  check  plats  in  the  same  series. 

Varietal  Variation. — Variation  in  protein  content  occurs  be- 
tween different  varieties  of  wheat,  even  though  the  strain  be  a 
pure  one  and  the  soil  conditions  under  which  they  are  grown  be 
the  same.  The  analyses  given  below  are  from  plants  of  pure 
strain  grown  during  the  course  of  breeding  experiments  in  cent- 
gener  plats  under  just  as  nearly  the  same  conditions  as  it  is 
possible  to  secure  in  the  field.  The  soil  was  uniform  and  the 
plants  were  4  inches  apart  each  way  and  seeded  at  a  uniform 
depth  of  2  inches. 

Grown  at  Davis,  1908 


Little  Club 

Number  of 
samples 

163 

Per  cent 
protein 

11.83 

Propo 
Fretes 

38 
111 

12.20 
13.46 

Grown  at 

Ceres,  1908 

Chul 

Number  of 
samples 

53 

Per  cent 
protein 

14.98 

Kharkov 

50 

13.95 

White  Australian 

70 

14.69 

In  addition  to  the  varietal  variations  here  shown,  the  above 
figures  are  of  interest  in  showing  that  Kharkov,  which  is  a  hard 
winter  wheat,  when  grown  under  California  conditions  in  the 
same  season  does  not  carry  a  higher  percent  of  protein  than  a 
good  quality  of  the  White  Australian  belonging  to  the  white 
wheat  class. 


1913]     Shaw:  Studies  on  Tnfuences  Affecting  Protein  Content  of  Wheat        67 

y aviation  in  Individual  Plants. — That  there  is  a  wide  dif- 
ference in  the  ability  of  individual  plants  to  elaborate  protein 
in  the  grain  is  shown  from  the  analysis  of  a  number  of  varieties 
grown  under  like  conditions  on  the  same  character  of  soils.  These 
plants  were  all  grown  in  the  same  season  in  centgener  plats,  the 
plants  being  four  (4)  inches  apart  each  way,  and  seeded  at  uni- 
form depth  by  means  of  a  centgener  planter.  The  outside  rows 
were  all  cut  away  at  harvest  and  discarded,  so  that  none  of  the 
plants  shown  here  represent  outside  rows,  but  all  had  a  uniform 
feeding  area  on  uniform  land. 

In  each  of  the  lots  shown  in  Table  1  it  will  be  noted  that  there 
is  a  great  variation  between  individual  plants  even  when  grown 
under  exactly  the  same  climatic  and  cultural  conditions.  The  vari- 
ation in  individual  plants  of  the  variety  White  Australian  ranged 
from  a  minimum  of  9.06  to  15.31  per  cent  total  protein,  or  a 
variation  of  6.25  per  cent  within  25  plants,  and  the  range  in 
Little  Club  is  even  greater,  being  from  7.12  to  15.22. 

Such  evidence  as  that  presented  above  would  seem  to  throw 
the  main  factors  determining  the  protein  content  of  wheats,  and 
inferentially  other  grains,  externally,  primarily  upon  some  cli- 
matic influence,  and  secondarily  upon  the  internal  factors  of 
variety  and  individuality  of  the  plant  itself.  It  seems  certain 
that  the  individuality  of  the  wheat  plant  is  just  as  potent  in 
determining  the  protein  content  of  the  grain  as  is  the  indiv- 
iduality of  the  dairy  cow  in  determining  the  fat  content  of  her 
milk  and  that  it  is  just  as  impossible  to  feed  protein  into  wheat 
by  increasing  the  available  nitrogen  of  the  soil  as  it  is  to  feed 
butter-fat  into  a  cow's  milk. 

Following  these  general  considerations  of  some  of  the  external 
and  internal  factors  bearing  upon  the  question  of  protein  vari- 
ation, attention  is  turned  to  more  restricted  phases  of  the  problem. 


68  University  of  California  Publications  in  Agricultural  Sciences    [Vol. 


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1913]   Shmv :  Studies  on  Influences  Afecting  Protein  Content  of  Wheat        69 

THE  CONSTANCY  OF  CHANGE  TOWARD  STARCHINESS  IN  WHEAT 

KERNELS 

The  phase  presented  in  this  portion  of  the  study  has  to  do 
with  the  tendency  of  varieties  toward  a  constancy  of  change 
from  the  type  seeded.  The  central  idea  was  that  if  there  really 
existed  a  strong  tendency  toward  the  development  of  a  distinctl}^ 
starchy  nature,  under  the  conditions  which  here  obtain,  not  only 
would  the  progeny  of  perfectly  typical  glutenous  kernels  show 
a  considerable  number  of  more  or  less  starchy  kernels  and  a 
lower  protein  (gluten)  content,  but  spotted  kernels  would  show 
a  similar  change  in  their  progeny ;  and  the  entirely  changed 
kernels,  carrying  100  per  cent  of  starchy  kernels  in  the  original, 
would  show  a  still  lower  percent  of  typical  kernels  and  a  very 
low  protein  content  in  the  progenj^  remaining,  in  fact,  essentially 
starchy. 

The  plan  of  the  experiment  which  extended  over  the  seasons 
of  1906-08,  inclusive,  was  to  grow  a  number  of  varieties  of  wheat 
at  several  points  in  the  state,  the  physical  and  chemical  con- 
dition of  the  originals  used  being  first  determined,  and  to  follow 
the  changes  in  the  progeny  in  each  instance  during  the  season 
and  the  total  period  of  the  trials. 

The  experiments  were  conducted  in  the  fields  at  Modesto, 
Ceres,  and  Tulare  as  typical  of  San  Joaquin  Valley  conditions, 
and  at  Yuba  City  and  Davis  as  representative  of  Sacramento 
Valley  conditions.  In  order  to  follow  better  such  changes  as 
occurred,  by  the  eye  as  well  as  by  analysis,  such  varieties  of 
wheat  were  selected  as  had  typical  kernels  of  distinctive  ap- 
pearance. 

THE    NITROGEN    OF    TYPICAL    vs.    SPOTTED    WHEAT    KERNELS 
OF  THE  SAME  VARIETY 

At  the  outset  a  number  of  varieties  of  wheat  were  hand- 
separated  into  three  primary  lots  or  groups,  based  upon  the 
physical  appearance  of  the  kernels,  as  follows : 

Group  I,  consisting  entirely  of  perfectly  typical  (glutenous) 
kernels ; 

Group  II,  consisting  of  kernels  of  the  same  varieties  of  wheat 


70  University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 

which  were  of  an  entirely  starchy  character,  showing  no  typical 
kernels  whatsoever; 

Group  III,  consisting  of  kernels  of  the  same  varieties  of  an 
intermediate  appearance  upon  which  the  percentage  of  typical 
kernels  was  determined  in  each  case. 

Each  of  these  lots  was  subjected  to  a  determination  of 
the  nitrogen  content  in  order  to  ascertain  whether  this  dis- 
tinctive appearance  of  the  kernels  was  a  fair  indication  of  their 
nitrogen  content. 

The  results  are  subtended: 

Nitrogen  Content  (Dry  Basis) 


Sample 

no. 

1 

Typical 
kernels 

1.75 

More  or 

less  starchy 

kernels 

1.45 

Sample 

no. 

2 

1.75 

1.54 

Sample 

no. 

3 

1.67 

1.54 

Sample 

no. 

4 

1.85 

1.57 

Sample 

no. 

5 

1.98 

1.74 

Sample 

no. 

6 

2.13 

"1.78 

Sample 

no. 

7 

2.09 

1.73 

Sample 

no. 

8 

2.03 

1.74 

Sample 

no. 

9 

1.98 

1.52 

Sample 

no. 

10 

2.02 

1.90 

Sample 

no. 

11 

2.12 

1.90 

Sample 

no. 

12 

2.27 

2.08 

Sample 

no. 

13 

1.82 

1.68 

Sample 

no. 

14 

2.03 

1.68 

Sample 

no. 

15 

2.33 

1.83 

Sample 

no. 

16 

1.83 

1.62 

Average  1.97  1.69 

These  results  seem  to  show  that  the  physical  appearance  of 
the  grain  may,  in  general  at  least,  be  taken  as  a  fair  index  of 
its  relative  gluten  or  protein  content. 

Each  of  the  above  described  groups  was  drill-seeded  in  a 
num})er  of  rows  in  a  uniform  manner  and  under  uniform  soil 
conditions  and  given  the  same  cultural  treatment. 

The  results  of  the  experiments  in  1906  are  tabulated  below: 


1913]     Shaw :  Studies  on  Influences  Affecting  Protein  Content  of  Wheat 


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1913]     Shaiv :  Studies  on  Influences  Afecting  Protein  Content  of  Wheat         73 

DISCUSSION  OF  1906  RESULTS 

Examination  of  the  1905-06  results  shows  that  in  physical 
appearance  none  of  the  seven  originals  carrying  100  per  cent  of 
typical  kernels  (Group  I)  maintained  their  perfection  in  their 
progeny — all  showed  some  starchy  kernels.  The  fact  that  those 
grown  at  Modesto  show^ed  97.5  per  cent  of  typical  kernels  in 
their  progeny  while  those  grown  at  Yuba  City  showed  but  60.1  per 
cent  ver}^  strongly  suggests  there  must  have  been  some  local 
condition  of  either  soil  or  climate  which  influenced  this  change. 

The  average  protein  content  of  this  group  at  Modesto  and 
Yuba  City  decreased  slightly,  as  did  the  percentage  of  typical 
kernels,  showing  13.15  per  cent  total  protein  in  the  originals  and 

12.91  per  cent  in  the  progeny ;  while  at  Yuba  City  the  results 
were  12.52  per  cent  for  the  originals  and  11.64  for  the  progeny. 
It  will  be  noted  also  that  the  Modesto  samples  possessed  some- 
what higher  total  protein  as  well  as  a  larger  percentage  of  typical 
kernels  than  did  those  grown  at  Yuba  City. 

Under  Group  II  at  Modesto  the  progeny  showed  an  average 
of  92.5  per  cent  of  typical  kernels  from  originals  carrying  abso- 
lutely none,  and  a  comparative  protein  content  of  10.97  per  cent 
for  the  originals  against  13.55  per  cent  for  the  progeny,  or  an 
increase  of  2.58  per  cent. 

Group  III  at  Modesto  carried  an  average  of  40.7  per  cent 
typical  kernels  in  the  originals  and  96.6  per  cent  in  the  progeny, 
and  a  protein  content  of  12.55  and  13.11  respectively,  a  distinct 
increase  in  both  respects.  At  Yuba  City  this  group  showed  62.9 
and  69.5  per  cent  typical  kernels  in  the  originals  and  progeny 
respectively,   and  a  protein  content   of   12.07   per   cent  against 

11.92  per  cent. 

Considered  as  a  whole  the  results  are  expressed  below: 


74 


University  of  California  Fublications  in  Agricultural  Sciences    [Vol.  1 


1906 
Original 
Progeny 


Group   I 


Number      Per  cent 
of  samples     typical 
in  group      kernels 


100.0 
81.5 


Per  cent 

total 
protein 

12.74 

12.78 


Group   II 


Group   III 


Number    Per  cent  Per  cent  Number    Per  cent  Per  cent 

of  samples  typical            total  of  samples    typical  total 

in  group     kernels  protein  in  group     kernels  protein 

9  0.0 


9 


75. 


10.58 
12.50 


55.6 

78.5 


12.23 
12.32 


General  General 

average  average 

Total         per  cent  per  cent 

number        typical  total 


1906 

of  trials 

kernels 

protein 

Original  

22 

47.39 

11.71 

Progeny  

22 

78.2 

12.54 

.  It  is  here  shown  that  the  average  per  cent  of  typical  kernels 
in  all  the  originals  was  47.4  and  in  the  progeny  78.2,  and  the 
protein  content  11.71  per  cent  and  12.54  per  cent  respectively. 
It  is  perfectly  evident  from  these  figures  that  a  wheat  which 
consists  of  100  per  cent  starchy  kernels  may,  in  a  single  season, 
revert  to  practically  a  perfect  wheat,  or  to  essentially  the  same 
average  protein  content  for  the  season  and  region  as  a  wheat 
groivn  from  perfectly  glutenous  kernels. 

Instances  of  this  are  particularly  shown  in  samples  no.  713 
and  no.  725.  In  the  first  the  original  consisted  of  100  per  cent 
of  entirely  starchy  kernels  and  still  produced  96.9  per  cent  of 
typical  kernels  in  the  progeny.  In  the  second  the  same  is  shown 
to  a  somewhat  smaller  extent,  the  original  carrying  but  38.3 
per  cent  typical  kernels  and  still  producing  97.9  per  cent  in  the 
progeny. 

Other  trials. — Trials  were  also  made  to  determine  the  per- 
centage of  typical  seed  in  six  varieties  grown  at  Modesto  from 
seed  in  which  the  percentage  of  typical  kernels  had  been  pre- 
viously determined,  using  1000  kernels  as  a  basis.  These  were 
also  seeded  in  rows  and  were  harvested  in  the  hard  dough  stage. 
Counts  were  then  made  to  determine  the  percentage  of  typical 
kernels  in  the  progeny.     The  results  appear  below. 


1913]     Shaw :  Studies  on  Influences  Afecting  Protein  Content  of  Wheat        75 


Name 

Per  cent 
of  typical 

kernels 

in  original 

1905 

Per  cent 

of  typical 

kernels 

1906 

Fretes 

94.0 

95.2 

Eedwinter 

51.0 

91.5 

Hungarian 

78.7 

95.3 

Kubanka 

38.3 

72.6 

Koola 

16.8 

93.9 

Marouani 

43.2 

93.1 

Average  53.7  90.2 

In  each  of  the  above  cases  the  percentage  of  typical  kernels 
increased  in  the  1906  crop  over  those  of  the  1905  crop  and  the 
average  increase  was  from  53.7  to  90.2;  it  seems  probable  that 
some  climatic  or  soil  factor  is  more  than  likely  to  have  been  the 
cause  of  this  change.  This  is  particularly  noticeable  upon  com- 
paring the  results  between  the  two  stations. 

The  above  figures  not  only  do  not  indicate  that  there  was 
any  strongly  marked  tendency  toward  lowering  in  quality,  but, 
on  the  other  hand,  the  general  tendency  seems  to  be  upward  for 
this  season,  particularly  at  Modesto. 

The  Relation  of  Moisture. — It  has  been  shown  repeatedly 
by  many  investigators  that  the  composition  of  plants  varies 
considerably  in  different  localities  and  in  different  seasons, 
and  that  the  principal  factor  seems  to  be  the  climate  instead 
of  the  soil,  variations  due  to  the  latter  being  very  slight  if 
any.  This  feature  will  be  presented  more  in  detail  later  in 
this  paper.  While  the  relation  of  the  water  content  of 
the  soil  at  various  stages  of  the  plant  growth  to  the 
protein  content  of  wheats  also  forms  a  portion  of  these  investi- 
gations for  later  presentation,  it  may  be  said  that  the  Utah 
Station  has  conducted  a  number  of  experiments  upon  the  effect 
of  water  upon  the  composition  of  plants,  the  method  of  which 
in  each  case  has  been  to  apply  different  amounts  of  water  through- 
out the  season  on  contiguous  plats  of  uniform  land.  The  fol- 
lowing selected  results  from  the  Utah  experiments  as  affecting 
the  protein  content  of  the  grain  are  of  interest  in  this  connection. 


76  University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 


Inches 

of 
water 
applied 

7.5 

Protein  content  of 

Corn 
kernels 

15.08 

Oat 

kernels 

20.79 

Wheat 
kernels 

26.72 

15.0 

13.48 

17.29 

19.99 

37.3 

12.52 

15.49 

16.99 

It  is  shown  here  that  as  the  amount  of  water  is  increased  the 
protein  content  decreases. 

Others  have  also  observed  the  effect  of  water  upon  the  com- 
position of  crops.  Mayer,  in  Holland,  showed  that  on  a  soil 
having  10  per  cent  of  water  the  crop  contained  10.6  per  cent  of 
protein,  while  on  a  soil  with  30  per  cent  of  water  the  protein 
percentage  was  only  6.6.  Carleton  calls  attention  to  the  fact 
that  in  the  same  varieties  of  wheat  grown  in  the  humid  and  arid 
regions  of  the  United  States  the  protein  content  was  11.94  per 
cent  for  the  former  and  14.4  per  cent  for  the  latter.  Experi- 
ments conducted  at  Rothamstead,  England,  show  that  barley  in 
a  wet  year  contained  9.81  per  cent  of  protein  and  in  a  dry  year 
12.99  per  cent. 

The  particular  connection  of  this  review  of  the  observation 
of  other  workers  to  the  results  here  presented  lies  in  the  dif- 
ferences in  the  results  secured  in  1906  between  the  Modesto  and 
the  Yuba  City  stations,  at  the  former  of  which  both  the  per- 
centage of  typical  kernels  and  the  protein  content  in  the  progeny 
was  distinctly  higher  than  at  the  latter.  These  three  factors  are 
contrasted  below. 

Inches  Per  cent  Per  cent 

rainfall  tj'pical  total 

Dec.  1-June  30  kernels  protein 

Modesto  11.94  95.35  13.50 

Yuba  City  26.68  63.99  11.74 

A  very  casual  observation  of  these  results  shows  that  both 
the  distinct  difference  in  percentage  of  typical  kernels  and  that 
between  the  protein  content  of  the  grain  of  the  two  stations  was 
without  doubt  very  strongly,  if  not  entirely,  determined  by  the 
difference  in  moisture  which  the  grain  received  after  planting. 

EXPERIMENTS  OF  1907 
In    1907   the   experiment  described   above  was   continued   at 
IModcsto   and    Yuba    City,    and   also   extended   to   Tulare.      The 
results  are  stated  below. 


1913  ]     Shmv :  Studies  on  Influences  Affecting  Protein  Content  of  Wheat 


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CDCiCiCiCiCiCiCiCiOiCiCiCiCiOiCiCiCiCiCiCiCiCiCiCiCi 


78  University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 


Oq^(Mr-l(MO]CvI(M(M<MC<l<M 


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1913]     Shaiv :  Studies  on  In^ucnces  Ajfecting  Protein  Content  of  Wheat        79 


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80 


Vniversitif  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 


Discussion  of  1907  Results.- — Essentially  the  same  results 
are  shown  here  as  in  the  preceding  season.  Of  the  seven  (7) 
originals  in  Group  I,  all  of  the  progeny  dropped  distinctly 
below  the  original  standard,  showing  55.99  per  cent  of 
typical  kernels,  while  the  eight  samples  in  Group  II,  carrying 
distinctly  starch  kernels  in  the  originals,  all  increased  by  about 
the  same  amount  as  the  others  dropped,  the  average  per  cent  of 
typical  kernels  in  the  progeny  being  55.2  per  cent,  or  less  than 
0.7  per  cent  below  those  of  Group  I. 

In  the  group  of  typical  kernel  originals,  only  one  out  of  seven 
increased  in  protein  content,  the  average  per  cent  in  the  originals 
being  12.79  against  11.20  per  cent  in  the  progeny,  or  a  drop  of 
1.59  per  cent.  In  Group  II,  consisting  entirely  of  the  distinctly 
starchy  kernels,  there  was  a  universal  increase  in  the  typical 
kernels  in  the  progeny,  and  three  out  of  eight  increased  in  protein 
content,  the  averages  being  10.65  per  cent  for  the  originals  and 
11.02  per  cent  for  the  progeny.  Group  III  showed  58.1  per 
cent  typical  kernels  in  the  originals  and  64.1  per  cent  in  the 
progeny,  an  increase  of  6  per  cent;  and  11.50  and  12.25  per 
cent  total  protein,  respectively. 

Summarizing  by  groups  as  in  the  results  of  the  preceding 
year,  the  figures  appear  as  below : 


Group   I 

Group   II 

Group   III 

1 

1907 

Number 

in 

group 

Per  cent 
typical 
kernels 

Per  cent 

total 
protein 

Number 

in 

group 

Per  cent 
typical 
kernels 

Per  cent 

total 
protein 

Number 

in 

group 

Per  cent 
typical 
kernels 

Per  cent                     , 
total                      4 
protein 

Original  . 

....     7 

100.0 

12.79 

8 

0.0 

10.65 

18 

58.10 

11.50 

Progeny  . 

7 

55.9 

11.24 

8 

55.2 

11.02 

18 

64.10 

12.25 

1907 
Original 
Progeny 


General 

average  General 

Total        per  cent  average 

number       typical  per  cent 

trials         kernels  protein 


33 
33 


58.1 
60.2 


11.56 
11.72 


1913  J     Shaw :  Studies  on  Influences  Affecting  Protein  Content  of  Wheat        81 

Eighteen  of  these  thirty-three  samples  showed  an  increase 
of  typical  kernels  in  the  progeny  over  the  originals,  the  respective 
average  being  60.2  and  54.6.  The  protein  content  increased  in 
sixteen  (16)  out  of  thirty-three  cases,  with  averages  for  originals 
and  progeny  of  11.56  per  cent  against  11.72  per  cent. 

EXPERIMENTS  OF  1908 

The  same  experiments  were  continued  in  1908  at  Davis  instead 
of  Yuba  City,  and  at  Ceres  instead  of  Modesto.  The  tabulated 
results  of  this  season  are  presented  below: 


82 


University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 


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1913]     Shaiv :  Studies  on  Influences  Afecting  Protein  Content  of  Wheat        83 


t^   rH   GO   O 

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84  University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 


Summarizing  the  results  of  the  foregoing  table  for  the  three 
groups  we  find  the  following : 


Group   I 

Group   II 

Group   III 

1908 

Number 

in 

group 

Per  cent 
typical 
kernels 

Per  cent 

total 
protein 

Number 

in 

group 

Per  cent 
typical 
kernels 

Per  cent 

total 
protein 

Number 

in 

group 

Per  cent 
typical 
kernels 

Per  cent 

total 
protein 

Original   . 

7 

100.0 

13.44 

Non 

e  of  this  type 

10 

72.23 

13.59 

Progeny   . 

7 

98.7 

15.14 

seeded 

10 

97.80 

14.54 

1908 

Total 

numbei 

trials 

General 
average 
per  cent 
typical 
kernels 

General 
average 
per  cent 
protein 

Original  

17 

83.6 

13.54 

Progeny  

17 

98.1 

14.78 

Discussion  of  1908  Results. — Out  of  a  total  of  seventeen  cases 
twelve  show  both  an  increased  number  of  typical  kernels  and 
an  increased  protein  content  in  the  progeny  over  the  originals, 
the  averages  being  as  follows : 

Original  Progeny 

Average  per  cent  typical  kernels 83.6  98.10 

Average  per  cent  total  protein 13.54  14.78 

In  Group  I,  three,  out  of  a  total  of  seven,  show  a  decrease  in 
the  percentage  of  typical  kernels  and  the  other  four  show  the 
same  as  the  original,  the  average  being  98.7  per  cent  with  an 
average  protein  content  of  13.49  per  cent  in  the  original  against 
15.14  in  the  progeny. 

In  Group  III,  consisting  of  ten  samples,  nine  show  an  increase 
in  both  percentage  of  typical  kernels  and  total  protein,  the 
average  being: 

Original  Progeny 

Average  per  cent  typical  samples  72.23  97.80 

Average  per  cent  total  protein  13.59  14.54 


The  results  expressed  by  groups  are  shown  in  the  following 
table : 


Number 

of 

trials 

21 

Per  cent  of  typical 
kernels 

A 

Per  . 
P 

;ent  total 
rotein 

A 

Group  I 

Original 
100.0 

Progeny 

78.7 

Original 
13.00 

Progeny 
13.04 

Group  II 

17 

0.0 

65.4 

10.61 

11.76 

Grou})    III 

34 

61.97 

80.1 

12.44 

13.03 

1913]     Shaw:  Studies  on  Influences  Affecting  Protein  Content  of  Wheat        85 

GENEEAL  DISCUSSION  AND  CONCLUSION 

Reviewing  the  results  of  the  three  years,  the  general  average 
of  typical  kernels  in  the  original  was  63.63  per  cent  and  of  the 
^eed  produced  therefrom  77.99  per  cent,  while  the  protein  con- 
tent was  12.34  per  cent  and  12.95  per  cent  respectively. 

The  results  as  a  whole  show: 

First — That  in  general  the  physical  appearance  of  durum  and 
red  wheats  is  a  fair  indication  of  their  relative  protein  content; 
kernels  having  a  distinctly  horny  or  glutenous  appearance  being 
higher  in  protein  than  those  of  a  more  or  less  dull  or  starchy 
appearance. 

Second — That  there  is  a  wide  seasonal  fluctuation  in  protein 
content  of  wheat  which  may  become  so  great  as  to  overbalance 
almost  entirely  any  hereditary  tendency  of  starchy  originals  to 
produce  the  same  characteristics  in  their  progeny. 

Third — That  the  protein  content  of  wheat  in  a  locality  is 
undoubtedly  largely  dependent  upon  the  seasonal  precipitation 
in  such  locality. 

Fourth — That  the  use  of  perfectly  typical  glutenous  seed  is 
invariably  followed  under  California  conditions  by  a  lowering 
of  the  gluten  content,  as  indicated  both  by  the  physical  appear- 
ance of  the  grain  and  by  its  protein  content. 

Fifth — That  if  the  original  carries  a  considerable  percentage 
of  starchy  kernels  the  progeny  usually  shows  an  increase  toward 
the  typical  character  to  a  degree  determined  by  the  character 
of  the  season  in  the  locality.  This  is  especially  so  with  reference 
to  the  precipitation,  which  is  in  some  instances  may  have  such 
a  strong  influence  as  to  cause  a  practically  perfect  grain  to  result 
from  an  original  seed  carrying  100  per  cent  of  starchy  kernels. 

Sixth — The  last  tabulation  by  groups  further  indicates  quite 
strongly,  however,  that  as  a  matter  of  fact  the  character  of  the 
seed  used  has  quite  a  marked  influence  upon  the  progeny,  and 
that  the  quality  of  the  seed  used,  to  some  degree  at  least,  deter- 
mines the  character  of  the  resultant  crop,  for  it  will  be  noted  that 
as  the  originals  decrease  in  both  percentage  of  typical  kernels 
and  protein  the  progeny  in  each  case  decrease  in  the  same  order, 
although  the  effect  of  this  is  materiallv  lessened  and  sometimes 


86 


University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 


almost   entirely   overcome   by   the   character   of   the   season,    as 
shown  by  the  other  results. 

THE  EELATION  OF  THE  TIME  OF  PLANTING  TO  THE  PROTEIN 
CONTENT  OF  WHEAT  KERNELS 

On  account  of  certain  differences  in  the  physical  appearance 
of  kernels  from  early  and  late  planted  wheats  in  preceding 
seasons,  experiments  were  conducted  in  1908  to  ascertain  more 
definitely  the  relation  of  early  and  late  planting  to  the  protein 
content  of  wheat  kernels.  In  these  experiments  it  was  intended 
to  conduct  parallel  series  of  plantings  at  the  University  Farm, 
Davis,  and  at  the  substation  at  Tulare.  At  the  former  place, 
however,  conditions  were  not  favorable  for  early  seeding,  and 
all  had  to  be  planted  too  late  to  be  at  all  satisfactory  in  pre- 
senting any  decided  contrast  in  planting  time,  particularly  for 
the  winter  wheats,  there  being  but  fifteen  days  between  the  early 
and  late  seeding.  The  first  plantings  of  the  season  were  not 
made  until  February  27,  and  the  second  plantings  on  March 
13.  At  Tulare  there  was  a  wider  difference,  the  plantings 
having  been  made  on  December  12,  and  January  17. 

As  a  matter  of  record,  however,  the  analyses  of  these  grains 
are  presented  below: 

Table  5. — Showing  the  Relation  of  Time  of  Planting  to  the  Physical 
AND  Chemical  Characteristics  of  Wheat  Kernels 


DAVIS,    1908 

First  ph 

mting 

Second  p' 

lanting 

Lab. 
No. 

2/05 

Typical 
kernels 
Per  cent 

100.0 

Total 
protein 
Per  cent 

11.81 

Gliadin 

Per  cent 

1.41 

Ash 
Per  cent 

1.76 

Typical 
kernels 
Per  cent 

67.1 

Total 
protein 
Per  cent 

10.67 

Gliadin 
Per  cent 

2.40 

Ash 
Per  cent 

2.04 

847A 

68.3 

11.19 

1.61 

1.76 

40.6 

10.62 

2.19 

2.03 

870A 

100.0 

13.78 

1.80 

1.73 

100.0 

13.74 

3.12 

1.72 

873A 

100.0 

13.38 

2.32 

1.73 

84.0 

13.08 

3.17 

1.72 

909A 

96.9 

14.41 

3.60 

2.07 

97.7 

14.89 

4.99 

1.49 

910A 

96.2 

12.99 

6.88 

1.93 

60.2 

9.77 

1.80 

2.03 

9111) 

95.6 

13.71 

4.16 

2.06 

99.4 

18.26 

8.05 

2.37 

91 7D 

100.0 

17.25 

6.95 

2.16 

100.0 

19.68 

5.41 

2.08 

91 9C 

85.7 

14.87 

5.88 

2.01 

99.4 

17.27 

5.82 

2.22 

922A 

99.0 

14.58 

4.15 

2.00 

99.1 

12.07 

3.09 

2.10 

595 

95.3 

14.91 

5.90 

1.89 

93.3 

13.24 

5.53 

2.13 

Av. 

94.2 

13.88 

4.06 

1.92 

85.5 

13.94 

4.14 

1.99 

1913]     Shaw :  Studies  on  In-jluences  Affecting  Protein  Content  of  Wheat        87 


TULARE, 

1908 

F'irst  planting 

Second  p^ 

lanting 

Lab. 
No. 

901 

Typical 
kernels 
Per  cent 

Total 
protein 
Per  cent 

10.34 

Gliadin 
Per  cent 

3.58 

Ash 
Per  cent 

Typical 
kernels 
Per  cent 

Total 
protein 
Per  cent 

11.30 

Gliadin 
Per  cent 

4.75 

Ash 
Per  cent 

896 

14.3 

9.82 

3.09 

1.92 

85.1 

12.95 

4.32 

1.68 

870 

20.7 

9.94 

3.43 

1.88 

68.1 

12.15 

4.97 

1.87 

848 

30.8 

10.96 

3.67 

2.03 

60.1 

11.53 

4.04 

1.84 

892 

74.5 

11.41 

3.51 

2.04 

98.1 

13.63 

4.48 

2.04 

338A6 

78.3 

11.47 

3.60 

1.92 

97.4 

13.52 

4.44 

2.08 

Av. 

43.7 

10.65 

3.48 

1.96 

81.7 

12.51 

4.50 

1.90 

Grand 

av.  68.9 

12.26 

3.77 

1.94 

83.6 

13.22 

4.32 

1.95 

For  the  reason  indicated  above  the  plantings  made  at  Davis 
should  not  be  regarded  as  having  much  bearing  upon  the  question. 
Considering  the  plantings  made  at  Tulare,  however,  it  will  be 
noted  that  in  every  case  the  late  plantings  showed  both  a  higher 
percentage  of  typical  kernels  and  a  higher  protein  and  gliadin 
content  than  the  early  plantings. 

The  average  in  the  case  of  the  Tulare  samples  was : 


Number 

of 

trials 

Per  cent 
typical 
kernels 

Per  cent 

total 
protein 

Per  cent 
gliadin 

Early  plantings  .... 

5 

43.7 

10.65 

2.48 

Late  plantings  

....       5 

81.7 

12.51 

4.50 

Further  trials  were  made  during  the  season  of  1909-10  at 
Davis  and  Tulare.  The  planting  dates  at  these  stations  were  as 
follows : 


Early 

Late 

Davis 

February  27th 

March  13th 

Tulare 

December  26th 

January  25th 

88  University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 

Table  6  Shoaving  the  Eelation  of  Time  or  Planting  to  the  Physical 
AND  Chemical  Characteristics  of  Wheat  Kernels 


DAVIS,   1909 

First  planting 

Second  planting 

Lab. 
No. 

870B 

Typical 
kernels 
Per  cent 

98.5 

Total 
protein 
Per  cent 

15.77 

Gliadin 
Per  cent 

5.77 

Ash 
Per  cent 

1.75 

Typical 
kernels 
Per  cent 

99.1 

Total 
protein 
Per  cent 

15.46 

Gliadin 
Per  cent 

6.16 

Ash 
Per  cent 

1.90 

873B 

94.8 

16.14 

6.00 

1.82 

97.8 

15.86 

6.49 

1.88 

847C 

99.0 

15.46 

6.05 

1.73 

98.0 

16.01 

6.52 

2.13 

909B 

55.0 

17.62 

8.16 

1.99 

58.2 

18.02 

7.87 

2.75 

910B 

97.9 

16.66 

6.20 

1.79 

100.0 

16.82 

5.94 

1.98 

922B 

99.0 

18.34 

8.01 

1.80 

99.1 

18.42 

5.68 

1.89 

9]7E 

98.0 

18.42 

8.35 

2.21 

100.0 

20.02 

7.44 

2.45 

388a5 

100.0 

18.18 

7.71 

2.17 

100.0 

18.58 

6.92 

2.28 

875B 

100.0 

18.42 

8.18 

2.02 

100.0 

19.20 

8.20 

2.39 

911E 

16.82 

8.11 

1.79 

19.44 

9.14 

2.12 

912D 

100.0 

20.00 

8.88 

2.28 

100.0 

]8.66 

6.39 

2.48 

914D 

98.0 

17.62 

7.86 

2.03 

100.0 

17.62 

7.91 

2.45 

Av. 

94.5 

17.45 

7.44 

1.94 

95.6 

17.84 

7.05 

2.22 

Table  6 — (Continued) 


Lab. 
No. 

1043aG 

]040aG 

]041aG 

]042aG 

1040aS 

]041aS 

]042aS 

1 043aS 

]151 

1155 

Av. 
Grand  a 


Typical 
kernels 
Per  cent 

77.0 

48.0 

44.0 

89.1 

35.0 

66.0 

100.0 
53.6 
99.6 

100.0 

71.2 

V.  82.8 


TULARE,    1909 


F'irst  planting 


Total 
protein 
Per  cent 

10.73 

11.13 

11.07 

13.12 

11.36 

10.50 

12.89 

10.62 

15.38 

17.20 

12.40 

14.92 


Gliadin 


Ash 


Per  cent    Per  cent 


3.45 
3.75 
3.79 
4.99 
4.11 
3.68 
5.06 
3.96 
7.32 
6.95 
4.71 
6.07 


2.23 
2.24 
2.14 


2.29 
2.10 
2.06 
2.37 


1.81 
2.15 
2.04 


Typical 
kernels 
Per  cent 

100.0 

100.0 

99.0 


100.0 
100.0 

100.0 
99.2 

100.0 
99.8 
97.7 


Second  planting 


Total 
protein 
Per  cent 

17.26 

17.21 

15.90 

18.91 

13.73 

10.33 

16.47 

16.47 

15.34 

17.24 

15.88 

16.86 


Gliadin         Ash 
Per  cent    Per  cent 


6.31 
5.50 
5.35 
6.65 
5.91 


6.19 


6.50 
5.38 
5.97 
6.51 


2.86 
2.20 
2.40 
2.59 
2.22 
2.34 
2.54 
2.74 


2.25 
2.46 
2.34 


These  results  show  that  out  of  twenty-two  (22)  cases,  seven- 
teen (17)  carried  a  higher  protein  content  in  the  later  plantings 
than  in  those  planted  early  and  that  a  like  number  out  of  the 
twenty-two  carried  either  a  larger  percentage  or  an  equal  number 
of  typical  kernels  in  the  late  plantings. 

The  averages  are  shown  below: 


1913]     Shaw :  Studies  on  Influences  A  f  eating  Protein  Content  of  Wheat        89 


Typical 

Total 

Trials 

kernels 

protein 

Gliadin 

Early  plantings  .... 

....     22 

82.8 

14.92 

6.07 

Late  plantings  

....     22 

97.7 

16.86 

6.51 

To  further  secure  data  upon  the  effect  of  the  time  of  planting 
upon  the  quality  of  the  grain,  twelve  (12)  varieties  of  wheat 
were  seeded  at  Ceres  on  four  different  dates,  in  rows  I6I/2  feet 
long  and  12  inches  apart,  as  follows : 

First  planting  November  28,  1908 

Second  planting  December  12,  1908 

Third  planting  December  31,  1908 

Fourth  planting  January  19,  1909 

The  grain  made  a  good  stand  and  grew  well  during  the  season 
and  matured  as  shown  in  the  following  table : 


Date  of 

ElPENING 

Lab. 

No. 

Date  of 

plantings 

Nov. 

28 

Dec. 

12 

Dec. 

31 

Jan. 19 

914 

June 

15 

June 

18 

June 

22 

July  17 

909 

June 

14 

June 

18 

June 

20 

July     7 

910 

June 

18 

June 

'20 

June 

20 

June  24 

911 

June 

16 

June 

20 

June 

26 

July     7 

915 

June 

20 

June 

24 

July 

3 

July     7 

913 

June 

16 

June 

20 

June 

24 

July     7 

917 

June 

15 

June 

16 

June 

19 

July     7 

916 

June 

11 

June 

18 

June 

22 

July     7 

919 

June 

16 

June 

20 

June 

24 

July     7 

912 

June 

16 

June 

18 

June 

22 

July     7 

1895 

June 

18 

June 

20 

June 

22 

June  24 

2/05 

June 

15 

June 

15 

June 

15 

June  15 

This  series  of  plantings  is  of  special  interest  in  that  out  of 
the  twelve  varieties  planted  all  but  one  show  the  highest  protein 
content  in  the  latest  planting,  and  the  one  exception  shows  the 
highest  in  the  third  planting.  In  this  case  the  difference  between 
the  protein  content  of  the  third  and  fourth  plantings  is  less  than 
one-half  of  1  per  cent.  It  is  further  of  interest  to  note  that 
there  was  quite  a  regular  increase  in  the  protein  content  in  the 
order  of  planting.  While  the  season  was  evidently  one  con- 
ducive to  the  development  of  a  relatively  high  quality  of  grain 


90  University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 

generally,  it  is  noticeable  that  the  average  percentage  of  typical 
kernels  also  follows  the  same  order  as  the  protein  content,  al- 
though the  difference  is  slight.  If  the  average  be  made  after 
discarding  the  last  sample,  which  constitutes  the  single  exception 
above  referred  to,  the  order  in  the  average  of  typical  kernels 
will  appear  absolutely  the  same  as  that  of  the  protein  content, 
and  very  consistently  bears  out  the  results  secured  at  Davis  and 
Tulare,  previously  discussed.  While  it  may  reduce  the  yield, 
relatively  late  seeding  tends  to  produce  a  grain  of  better  quality 
than  does  early  planting. 

This  same  fact  is  further  evidenced  by  a  series  of  thirty- 
seven  (37)  types  of  New  South  Wales  hybrid  stocfe  seeded  at 
Davis  in  the  season  of  1910-11 ;  the  dates  of  planting  being 
December  12,  1910,  and  February  20,  1911.  The  analyses  from 
this  lot  are  tabulated  below : 


1913]     Shaiv :  Studies  on  Influences  Affecting  Protein  Content  of  Wheat        91 


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92  Vniversity  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 

Table  8  Showing  the  Relation  of  Time  of  Planting  to  the  Gluten 
Content  of  Wheat  Kernels 


Early  plantin 

g 

Late  planting 

Sample 
No. 

Total 
gluten 

Total 
gliadin 

Ash 

Total 
gluten 

Total 
gliadin 

Ash 

1 

14.37 

5.890 

1.71 

16.41 

6.742 

1.88 

2-1 

13.00 

5.322 

1.71 

13.42 

5.787 

1.78 

3 

13.74 

5.552 

1.70 

14.54 

5.839 

2.05 

4-B 

14.42 

5.657 

1.73 

15.22 

6.049 

1.76 

4-C 

12.66 

5.555 

1.63 

15.44 

6.503 

1.69 

4-D 

12.15 

4.572 

1.48 

15.22 

5.594 

1.86 

4-E 

10.79 

4.646 

1.82 

13.46 

6.299 

1.96 

5-2 

11.98 

4.311 

1.78 

15.28 

5.248 

1.73 

5-4 

12.55 

4.186 

1.81 

16.41 

6.477 

1.61 

5-5 

12.38 

4.657 

1.88 

15.22 

5.935 

1.61 

6 

11.26 

4.287 

1.91 

16.30 

5.873 

1.94 

7 

13.80 

5.424 

1.94 

17.38 

6.844 

1.83 

8 

14.42 

6.179 

1.74 

16.24 

7.242 

1.60 

10-1 

13.46 

4.595 

1.77 

16.01 

5.600 

1.70 

10-2 

12.21 

3.765 

1.78 

15.33 

4.975 

1.71 

10-3 

13.06 

4.333 

1.61 

15.50 

5.731 

1.73 

10-4 

11.92 

3.845 

1.75 

16.01 

5.560 

1.81 

11 

12.62 

5.210 

1.74 

14.94 

6.117 

1.97 

12-2 

14.25 

5.339 

1.76 

14.59 

5.765 

1.79 

12-4 

13.06 

4.731 

1.70 

14.93 

5.288 

1.51 

13 

13.57 

4.884 

1.70 

14.94 

5.310 

1.77 

15 

13.34 

4.879 

1.81 

14.14 

5.026 

1.95 

18 

14.02 

4.311 

1.64 

16.13 

5.469 

1.63 

20 

13.51 

5.418 

1.72 

15.56 

6.162 

1.60 

21 

11.70 

4.544 

1.56 

15.05 

5.077 

1.63 

22 

13.17 

4.328 

1.81 

16.41 

5.452 

1.58 

23 

16.01 

5.707 

2.05 

17.89 

6.747 

2.08 

24 

13.17 

4.708 

2.15 

15.44 

5.384 

1.80 

25 

14.08 

4.288 

2.10 

15.79 

5.305 

1.92 

26 

13.34 

4.461 

1.82 

15.84 

5.163 

2.04 

27 

14.08 

4.345 

1.87 

17.60 

6.088 

1.98 

28 

14.76 

5.117 

1.63 

15.39 

5.350 

1.78 

29 

11.47 

3.311 

1.87 

14.54 

4.146 

1.76 

30 

13.24 

5.293 

1.65 

14.82 

5.867 

1.70 

31 

13.57 

5.572 

1.50 

14.54 

5.992 

1.94 

32 

13.26 

5.305 

1.58 

14.37 

6.054 

1.64 

33 

12.49 

4.515 

1.87 

14.54 

6.418 

1.81 

Av. 

13.158 

4.812 

1.764 

15.428 

5.796 

1.787 

In  this  series  of  plaritirios,  without  a  single  exception,  the  late 
planted  saini)les  r-nn  hij^her  in  protein  than  those  planted  early, 
and  the  same  holds  true  of  their  gliadin  content. 


1913]   Shaw:  Studies  on  Influences  Affecting  Protein  Content  of  Wheat        93 

General  Discussion 

Collecting  all  results  for  the  season  of  1908,  1909,  and  1911, 
the  following  grand  averages  appear  upon  which  to  base  con- 
clusions : 


No. 
trials 

17 

First  planting 

Second  planti 

ng 

Year 
1908 

Per  cent 
typical 
kernels 

78.5 

Per  cent 

total         Per  cent 
protein        gliadin 

12.75         3.84 

Per  cent 
typical 
kernels 

84.3 

Per  cent 

total 
protein 

13.45 

Per  cent 
gliadin 

4.26 

1909 

34 

87.9 

15.66         5.92 

98.4 

16.95 

6.39 

1911 

37 

13.16         4.84 

15.43 

5.80 

Av. 

88 

83.2 

13.85         4.87 

91.3 

15.28 

5.48 

These  figures  show  quite  definitely  that  late  planting:  tends 
toward  the  production  of  a  grain  carrying  a  higher  percentasre 
of  total  protein  and  gliadin,  as  well  as  a  higher  percentage  ol* 
typical  kernels,  than  does  early  planting.  Of  the  eighty-eight 
trials  made,  if  we  except  those  at  Davis  the  first  season  when 
the  two  plantings  were  made  at  dates  too  close  together  for  dis- 
tinctive results,  there  is  a  great  unanimit}^  of  results  in  favor 
of  the  late  seeded  grain  in  the  respects  indicated.  Considered 
as  a  whole,  the  late  plantings  show  an  increase  over  the  early 
plantings  of  8.1  per  cent  of  typical  kernels,  1.43  per  cent  of 
protein  and  .61  per  cent  of  gliadin.  Of  particular  note  are  the 
four  successive  plantings  made  at  Ceres  in  1909,  in  which  there 
is  a  regular  increase  in  protein,  gliadin,  and  typical  kernels  from 
the  early  to  the  latest  plantings,  and  also  the  plantings  at  Davis 
in  1910-11  in  which  the  same  unanimity  in  respect  to  protein 
and  gliadin  is  shown. 

The  results  covering  this  season  are  so  uniformly  in  favor 
of  the  late  planting  as  developing  the  higher  protein  and  gliadin 
content  that  the  question  seems  to  be  very  decisively  answered. 

THE  EFFECT  OF  THE  TIME  OF  HARVESTING  UPON  THE  PROTEIN 
CONTENTS  OF  WHEATS 

In  most  parts  of  the  country  from  which  is  obtained  the 
recognized  high  gluten  wheat  the  grain  is  cut  when  in  the  hard 
dough  stage  by  means  of  a  self-binder  and  later  threshed  by 
means  of  a  stationary  threshing  outfit.    In  California  practically 


94  University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 

all  of  the  grain  is  left  standing  on  the  straw  in  the  field  until 
it  is  ''dead  ripe"  and  finally  cut  in  that  condition  and  threshed  in 
a  single  operation  by  means  of  a  combined  harvester. 

In  the  later  method  the  grain  often  stands  on  the  straw  and 
is  subjected  to  the  action  of  the  sun  for  several  weeks  after 
reaching  the  hard  dough  stage.  This  difference  in  the  method 
of  harvesting  has  led  to  the  belief  among  many  that  the  time 
of  cutting  might  be  one  of  the  causes  of  the  relatively  low  protein 
(gluten)  content  of  Pacific  Coast  wheats. 

With  the  idea  of  determining  the  effect  of  such  standing  in 
the  field,  a  series  of  experiments  was  planned  to  ascertain  what 
might  be  the  influence  of  this  practice. 

For  the  purposes  of  this  experiment  several  varieties  of  wheat 
whose  kernels  possessed  distinctive  characteristics  as  indicated 
in  the  preceding  discussion  were  hand-separated  into  three  groups 
as  previously  shown  (page  69).  These  were  drill-seeded  under 
uniform  soil  and  climatic  condition  at  the  several  stations.  One- 
half  of  each  lot  was  harvested  in  the  hard  dough  stage,  in  which 
condition  it  would  normally  be  cut  by  a  binder,  and  the  other 
half  was  left  standing  on  the  straw  in  the  field  until  it  reached 
a  condition  suitable  to  be  cut  with  a  combined  harvester.  This 
was  finally  harvested  and  each  portion  subjected  to  analysis  in 
addition  to  determining  the  percentage  of  typical  kernels  as 
indicated  by  the  physical  appearance  of  the  kernels,  using  1000 
kernels  as  a  basis. 

Aside  from  the  main  question  as  to  the  effect  of  the  time  of 
cutting  upon  the  quality  of  the  grain,  the  data  as  presented  in 
tabular  form  further  furnishes  a  means  for  comparing  the 
original  used,  with  the  progeny  as  in  the  previously  discussed 
experiment  (pages  69-74). 


1913]     SJiaiv :  Studies  on  Influences  Afectmg  Protein  Content  of  Wheat 


95 


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96 


University  of  California  Publications  in  Agricultural  Sciences    [Vol. 


(M    C\r    CM    OCI    (M    (M 


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cococococococococococococococococococococococococococococococo 


1913]     Shaw :  Studies  on  Influences  Afecting  Protein  Content  of  Wheat        97 

Discussion  of  1906  Results. — In  these  results  the  same  thing 
is  noticeable  as  in  the  former  series,  viz.,  that  the  samples  grown- 
at  Modesto  as  a  whole  maintained  both  a  higher  percentage  of 
typical  kernels  and  a  higher  protein  content  than  those  grow^n 
at  Yuba  City,  the  comparison  by  averages  being  as  follows : 

Modesto         Yuba  City 
Average  per  cent  typical  kernels  89.70  65.2 

Average  per  cent  protein  11.87  10.5 

It  is  particularly  noticeable  that,  outside  of  the  instances  in 
which  the  originals  carried  100  per  cent  of  typical  kernels,  there 
was  in  this  season  a  practically  universal  marked  increase  in 
the  number  of  typical  kernels  in  the  progeny;  that  in  the  case 
of  sample  no.  714,  which  carried  no  typical  kernels,  the  original 
seed  being  100  per  cent  starchy,  the  second  cutting  produced 
100  per  cent  of  typical  kernels  and  the  first  cutting  (no.  713) 
96.9  per  cent  of  such  kernels.  Almost  as  good  a  showing  also  is 
made  bj^  samples  nos.  719  and  720,  which  produced  respectively 
96.1  and  96.3  typical  kernels  from  entirely  starchy  kernels. 

It  is  further  shown  that  in  the  case  of  the  early  cutting 
twenty-one  out  of  the  thirty-one  trials  show^ed  an  increase  in  the 
percentage  of  typical  kernels  in  the  progeny  over  the  original, 
the  relation  in  those  cases  in  which  increase  occurred  being  42.5 
per  cent  in  the  original  and  83.19  per  cent  in  the  progeny,  while 
in  the  eight  cases  in  which  decrease  occurred  the  averages  were 
96.1  per  cent  typical  kernels  in  the  originals  against  75.3  per  cent 
in  the  progeny,  which,  is  entirely  confirmatory  of  the  results 
presented  in  Table  2. 

Looking  more  parti cularW  at  the  specific  question  involved 
in  this  experiment,  viz.,  the  effect  of  the  time  of  cutting  upon 
the  percentage  of  typical  kernels  and  the  protein  content,  it  is 
found  that  the  following  averages  hold: 

Modesto 

Number       Per  cent 

of  typical  Per  cent 

trials  kernels  protein 

Early  cutting  15  94.6  13.38 

Late  cutting  15  90.1  13.19 


98  University  of  California  Fublicaiions  in  Agricultural  Sciences    [Vol.  1 


Yuba  City 

Number       Per  cent 

of  typical  Per  cent 

trials  kernels  protein 

Early  cutting  16  66.1  11.98 

Late  cutting  16  64.6  12.05 

These  average  figures  show,  in  the  case  of  the  Yuba  City  lot, 
an  apparent  contradiction  in  that  the  early  cutting  with  66.1 
per  cent  typical  kernels  showed  11.98  per  cent  protein,  while  the 
late  cutting  with  but  64.6  typical  kernels  showed  12.05  per  cent 
protein.  When  it  is  remembered,  however,  that  the  difference 
in  the  two  cuttings  is  represented  by  but  a  single  kernel  in  one 
hundred  and  that  the  average  protein  content  is  but  seven- 
hundredths  of  1  per  cent,  it  will  be  seen  that  the  discrepancy  is 
only  apparent,  and  well  within  the  limits  of  error  in  either  count 
or  analysis. 

As  a  whole,  it  is  shown  that  out  of  the  total  of  31  cases  11 
showed  an  increase  of  typical  kernels  in  the  late  cut  grain  and 
14  showed  an  increased  protein  content,  with  the  following  rel- 
ative averages: 

Number       Per  cent 

of  typical  Per  cent 

trials  kernels  protein 

Early  cuttings  31  79.89  12.66 

Late  cuttings  31  76.9  12.60 

The  results  for  1906  do  not  show  that  there  is  any  marked 
difference  to  be  attributed  to  allowing  the  grain  to  become  thor- 
oughly ripe  on  the  straw,  as  is  generally  done  in  California. 

EXPERIMENTS  OF  1907 

The  experiments  as  to  the  effect  of  the  time  of  cutting  wheat 
upon  its  protein  content  were  continued  in  the  manner  described 
above  in  the  season  of  1907  at  Modesto,  Yuba  City,  and  Tulare, 
with  the  following  results: 


1913]     Shaiv :  Studies  on  Influences  Afecting  Protein  Content  of  Wheat 


99 


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100         University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 


Cvii-!T-l(M(MC<l.-IC<J 


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1913]     Shaw :  Studies  on  Influences  Afecting  Protein  Content  of  Wheat      101 


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102  University  of  California  Fublications  in  Agricultural  Sciences    [Vol.  1 

The  averages  for  the  three  localities  are  shown  in  the  following 
small  table: 

Table  Showing  the  Average  Results  From  Early  and  Late  Cut  Wheats 
AT  Three  Station  in  1907 


Number 

of 

trials 

12 

Early  cut 

Late  cut 

A 

Modesto 

Per  cent 

typical         Per  cent 
kernels         protein 

52.3             9.92 

Per  cent 

typical         Per  cent 
kernels          protein 

49.2            10.40 

Yuba  City 

13 

73.0           11.62 

73.3           12.19 

Tulare 

5 

56.4           11.00 

57.8           10.92 

Grand  av. 

.     30 

62.6           10.80 

61.9           11.24 

From  a  total  of  30  cases,  12  showed  either  an  equal  or  larger 
per  cent  of  typical  kernels  in  the  late  cutting  than  in  the  early 
cutting,  a  number  altogether  too  large  to  indicate  that  the  late- 
ness of  cutting  had  any  material  influence  in  this  direction,  and 
this  is  further  shown  from  the  fact  that  the  general  average 
shows  62.6  per  cent  of  typical  kernels  in  the  early  cut  lots  and 
61.9  per  cent  in  the  late  cut  lots. 

Again  this  is  indicated  in  the  protein  content,  for  out  of  the 
30  cases  15  show  a  larger  percentage  in  the  late  cut  grain  than 
in  the  early  cut  lots,  and  the  general  average  in  protein  content 
is  essentially  the  same  in  the  two  lots,  viz.,  10.80  and  11.24  per 
cent  respectively. 

For  the  two  seasons  the  record  stands  as  follows : 


Number 

of 

trials 

27 

Early  cut 

Late  cut 

Modesto 

Per  cent 

typical         Per  cent 
kernels          protein 

77.68         11.8 

Per  cent 

typical         Per  cent 
kernels          protein 

73.7           11.95 

Yuba  City 

29 

69.1           11.96 

68.5           11.20 

Tulare 

5 

56.4           11.00 

57.8           10.92 

Average 

61 

71.1           11.80 

69.9           11.51 

These  figures  seem  to  show  quite  clearly  that  to  allow  the 
grain  to  stand  on  the  straw  until  in  the  proper  condition  for 
handling  with  the  combined  harvester  does  not  in  any  manner 
militate  against  its  (juality  either  in  physical  appearance  or  pro- 
tein content,  and  slioukl  set  at  rest  any  further  discussion  upon 
this   point. 


1913]     Shaw :  Studies  on  Injiuences  Affecting  Protein  Content  of  Wheat      103 

EFFECT  OF  SUNSHINE  ON  THE  COMPOSITION  OF  THE  WHEAT 

KERNEL 

It  has  long-  been  a  matter  of  common  knowledge  that  the 
composition  of  plants  varies  greatly  under  different  conditions 
and  in  different  localities.  The  exact  environmental  factor  or 
factors  which  cause  this  variation  in  composition  has  been  the 
cause  of  much  discussion  and  investigation. 

It  has  already  been  remarked  above  that  in  1882  Richardson 
observed  that  wheat  grown  in  Colorado  had  by  reputation  a 
much  higher  gluten  content  than  wheat  grown  from  the  same 
seed  in  Oregon.  Not  only  was  there  a  marked  difference  in  the 
composition  of  the  wheat  grown  in  the  two  states,  but  he  found 
that  the  wheat  grow^n  in  Colorado  had  a  higher  gluten  content 
than  the  original  seed,  while  that  grown  in  Oregon  had  a  lower 
gluten  content  than  the  seed  from  which  it  was  produced.  From 
these  observations  Richardson  and  Blount  concluded  that  the 
soil  was  the  modifying  factor. 

Wiley,  however,  draws  the  conclusion  that  the  difference  in 
gluten  content  is  due  to  climatic  conditions. 

Lawes  and  Gilbert  in  an  elaborate  series  of  experiments  have 
shown  that  the  use  of  manures  and  fertilizers  have  very  little 
influence  on  the  composition  of  the  wheat  kernel.  On  the  other 
hand,  they  found  a  wide  variation  in  composition  in  different 
seasons, 

Deherain  in  France  also  observed  that  a  difference  in  seasonal 
conditions,  especially  during  the  ripening  period,  had  a  marked 
influence  on  the  g'luten  content  of  wheat. 

Similar  observations  have  been  noted  by  Thatcher  of  Wash- 
ington, and  others. 

From  these  numerous  observations  and  experiments  it  has 
come  to  be  generally  conceded  that  the  gluten  content  is  influ- 
enced mostly,  if  not  wholly,  by  the  climate. 

Wheat  grown  in  the  coast  states  is,  as  a  class,  much  lower  in 
gluten  content  than  wheat  grown  in  the  central  west,  or  the 
northwestern  states.  Even  when  seed  of  a  high  gluten  content 
is  introduced,  a  product,  as  Richardson  observed,  considerably 
inferior  is  the  result. 


104  University  of  California  Puhlications  in  Agricultural  Sciences    [Vol.  1 

Climate  includes  a  large  number  of  factors.  The  specific 
climatic  factor  which  is  the  cause  of  this  variation  has  recently 
been  the  subject  of  much  study  and  investigation. 

The  formation  of  organic  compounds  in  the  plant,  such  as 
starch  and  gluten,  is  a  physiological  process.  The  maximum 
development  of  these  compounds  is  necessarily  dependent  upon 
favorable  conditions.  Starch  is  formed  under  the  influence  of 
sunlight.  The  formation  of  nitrogenous  compounds  requires  not 
only  an  adequate  supply  of  nitrogen,  but  also  a  supply  of  carbon 
compounds  in  the  proper  form  and  under  the  proper  conditions. 
Just  what  these  conditions  are  is  not  definitely  known. 

In  general,  investigators  have  concluded  that  the  large  amount 
of  sunshine  prevalent  in  the  coast  states  during  the  period  in 
which  the  seed  develops  works  directly  in  favor  of  the  formation 
of  large  amounts  of  starch.  In  other  words,  that  the  gluten 
content  is  low  only  by  reason  of  the  formation  of  proportionally 
larger  amounts  of  starch.  Were  this  the  case,  the  exclusion  of 
portions  of  the  sunlight  should  tend  to  increase  the  percentage 
of  gluten.  Theoretically  the  gluten  would  increase  inversely  with 
the  amount  of  sunlight  which  the  plants  received. 

In  order  to  determine  w^hether  or  not  sunlight  is  a  prominent 
factor  a  series  of  experiments  were  planned  in  which  portions 
of  the  natural  sunlight  were  excluded  from  the  growing  plants. 

Duplicate  experiments  were  conducted  during  the  seasons  of 
1908  and  1909  at  the  Tulare  Sub-station  and  at  the  University 
Farm  at  Davis. 

Original  Seed. — At  the  Tulare  Station  in  1908  the  originals 
used  were  as  follows : 

No.  864/07  Weissenberg 

No.  879/07  Currell 

No.  892/07  Beloglina 

No.  894/07  Velvet  Don  (starch  grains) 

No.  901/07  Turkey  Red  (gluten  grains) 

No.  899/07  Yellow  Gharnovka  (gluten  grains) 

At  Davis  the  following  varieties  were  used: 

No.  868/07  Kubanka 

No.  847  Kubanka 

No.  870A  Velvet  Don  (gluten  grains) 


1913]     Shaw :  Studies  on  Iniiuences  Affecting  Protein  Content  of  Wheat      105 


The  originals  showed  the  following  composition : 


At  Tulare 

Lab. 
No. 

Name 

Per  cent 
typical 
kernels 

Kernels 
in  10 
grams 

Per  cent 
protein 

Per  cent 
gliadin 

Per  cent 
ash 

864 

Weissenberg 

95.9 

325 

15.25 

3.63 

2.66 

879 

Currell 

87.6 

351 

13.55 

5.06 

2.11 

892 

Beloglina 

59.4 

271 

10.96 

3.84 

2.33 

894 

Velvet  Don 

0.0 

230 

8.79 

2.90 

2.00 

901 

Turkey  Eed 

100.0 

278 

13.41 

5.28 

1.99 

899 

Gharnovka 

100.0 

280 

12.20 

4.26 

2.02 

At  Davis 

Lab. 

No. 

Name 

Per  cent 
typical 
kernels 

Kernels 
in  10 
grams 

Per  cent 
protein 

Per  cent 
gliadin 

Per  cent 
ash 

868 

Kiibanka 

89.2 

184 

23.64 

4.67 

2  22 

847 

Kiibanka 

48.6 

11.03 

3.88 

1.97 

870-G 

Velvet  Don 

90.1 

239 

17.38 

4.80 

2.05 

Two  rows  of  each  of  these  varieties  were  drilled  parallel  into 
a  bed  24  feet  long.  In  the  spring  after  the  grain  began  to  make 
an  upright  growth  a  series  of  lath  screens  was  constructed  and 
placed  across  the  plat  so  as  to  shut  off  different  proportions  of 
the  direct  sunshine.  The  plat  was  thus  divided  into  sections. 
At  Tulare  sectional  screens  were  arranged  so  that  they  excluded 
three  quarters,  one  half,  and  one  quarter  of  the  sunshine  respect- 
ively. One  quarter  was  left  unshaded,  and  this  received  the  full 
sunshine.  In  this  manner  one  fourth  of  each  variety  received  the 
same  amount  of  sunshine.  At  Davis  a  similar  arrangement  was 
used,  except  that  the  amount  of  sunshine  admitted  was  one  third, 
one  half,  two  thirds,  and  full.  These  screens  were  left  in  place 
until  the  grain  was  matured  and  harvested.  Each  portion  of  each 
variety  thus  treated  was  harvested  separately  and  taken  to  the 
laboratory  for  analysis.     The  results  are  stated  below: 


106  University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 

Table  12. — Showing  Effect  of  Various  Amounts  of  Sunshine  on  the 
Gluten  Content  of  Wheat 


864 
894 
899 
901 
879 
892 
Av 


One-quarter 
sunshine 


Total 
gluten 

17.55 

12.10 

12.66 

15,33 

18.17 

15.39 

15.19 


Gliadin 
8.000 
4.930 
5.333 
6.602 
8.000 
6.674 
6.589 


TULARE,    1908 
One-half 
sunshine 


Total 
gluten 

18.00 

12.72 

12.61 

15.45 

18.63 

16.58 

15.665 


Gliadin 
8.065 
5.350 
5.311 
6.986 
8.406 
7.157 
6.879 


Three-quarters 
sunshine 


Total 
gluten 


Gliadi 


Full 
sunshine 

Total 
gluten 

Gliadin 

20.73 

9.599 

13.57 

4.856 

13.63 

5.515 

16.13 

6.545 

18.34 

8.463 

17.04 

7.554 

16.57 

7.088 

Gliadin  ratio      43.40 


43.91 


42.77 


DAVIS,    1908 


847 
870G 

Av, 


One-third 
sunshine 


Total 
gluten 

13.19 
12.69 
14.30 
13.39 


Gliadin 
4.17 
4.46 
4.71 
4.445 


Gliadin  ratio       33.19 


One-half 
sunshine 


Total 
gluten 

13.94 
12.37 


Gliadin 
4.17 
4.09 


13.15       4.13 
31.40 


Two-thirds 
sunshine 


Total 
gluten 

14.10 

13.40 

13.53 

13.67 


Gliadin 
4.42 
3.50 
4.21 
4.034 


Full 
sunshine 


Total 
gluten 

13.79 

12.47 

13.17 

13.14 


Gliadin 
4.28 
4.00 
4.52 
4.266 


29.57 


32.46 


EXPERIMENTS  OF  1909 
During  the  season  of  1909  the  same  experiment  was  conducted 
at  the  Tulare  and  Davis  Stations,  with  the  exception  that  the 
following  varieties  of  wheat  were  used: 


Gluten 
Gluten 
Starch 
Gluten 
Gluten 
Starch 


grains  from  Richi  1044A; 
grains  from  Kubanka  1045 A: 
grains  from  Kubanka  1045 A; 
grains  from  Turkey  1046A; 
grains  from  Beloglina  1047A 
grains  from  Beloglina  1047A, 


Determinations  made  upon  these  previous  to  seeding  showed 
as  follows : 


No. 
1044A-G 
1 045  A-G 
1045A-S 
1046A-G 
1047  A-G 
1047A-S 


Typical 

kernels 

Kernels 
in  10 
grams 

Protein 

Gliadin 

Ash 

]00.0 

189 

10.42 

3.17 

1.90 

100.0 

197 

13.09 

4.32 

1.74 

0.0 

220 

9.37 

2.59 

1.88 

100.0 

389 

15.43 

4.90 

2.21 

100.0 

372 

16.11 

5.33 

2.25 

0.0 

329 

7.37 

2.00 

1.77 

1913  ]     Shaw :  Studies  on  Influences  Affecting  Protein  Content  of  Wheat      107 


At  Davis  the  originals  and  their  analyses  were  as  follows 


No. 

Name 

Typical 
kernels 

Kernels 
in  10 
grams 

Protein 

Gliadin 

Ash 

870G 

Velvet  Don 

100.0 

227 

15.92 

4.27 

2.02 

870S 

Velvet  Don 

0.0 

239 

13.96 

4.73 

2.61 

1045A-G 

Kubanka 

100.0 

197 

13.09 

4.32 

1.74 

1045A-S 

Kubanka 

0.0 

220 

9.37 

2.59 

1.88 

1041A-G 

Kubanka 

100.0 

177 

13.05 

4.12 

2.02 

1041A-S 

Kubanka 

0.0 

194 

10.56 

2.99 

2.01 

The  gluten  and  gliadin  content  of  the  wheat  samples  from 
the  experiments  of  1909  are  reported  in  Table  13. 

Table  13. — Showing  Effect  of  Various  Amounts  of  Sunshine  on  the 
Protein  Content  of  Wheat 


Tulare,  1909 

One-quarter 
sunshine 

One-half 
sunshine 

Three-quarters 
sunshine 

Full 
sunshine 

1046G 

'  Total 
protein 

10.62 

Gliadin 
3.516 

Total 
protein 

7.38 

Gliadin 
5.043 

Total 
protein 

Gliadin 

Total 
protein 

14.77 

Gliadin 
5.651 

1047AS 

10.96 

3.998 

12.32 

4.413 

15.56 

5.577 

13.18 

4.084 

1047AS 

13.74 

4.737 

1045AS 

11.64 

4.100 

12.04 

4.271 

13.86 

3.783 

13.35 

3.839 

1045AG 

12.38 

3.902 

13.25 

4.674 

11.30 

4.312 

13.91 

4.981 

1044AG 

10.90 

3.271 

11.41 

3.436 

14.99 

4.385 

11.59 

3.720 

Av. 

11.30 

3.757 

11.27 

4.368 

13.92 

4.514 

13.42 

4.502 

Gliadin  ratio 

33.42 

38. 

74 

32.42 

33.54 

Davis,  1909 

One-third      . 
sunshine 

A 

One-half 
sunshine 

A 

Two 
sur 

-thirds 
ishine 

A 

Full 
sunshine 

1041AS 

Total 
protein 

15.34 

Gliadin 
4.845 

Total 
protein 

16.24 

Gliadin 
4.850 

Total 
protein 

16.41 

Gliadin 
5.151 

Total 
protein 

16.02 

Gliadin 
4.970 

1045AG 

14.71 

5.186 

14.37 

4.748 

15.50 

4.055 

14.48 

4.635 

1045AS 

14.71 

5.680 

* 

15.56 

6.248 

14.25 

5.850 

870AG 

16.53 

5.341 

* 

15.62 

4.867 

15.66 

5.214 

870AS 

15.79 

6.816 

14.06 

3.770 

15.50 

6.418 

15.50 

6.645 

1041AG 

16.36 

6.832 

16.70 

6.475 

17.09 

6.929 

16.30 

6.418 

Av. 

15.57 

5.773 

15.34 

4.960 

15.94 

5.611 

15.37 

5.622 

Gliadin  ratio 

36.82 

32.33 

35.20 

36.57 

*  There  was  not  enough  material  for  a  re-analysis  of  the  sample. 


108  University  of  California  Puhlications  in  Agricultural  Sciences    [Vol.  1 

Discussion. — If  the  proportion  of  protein  to  starch,  or  the  per- 
centage of  nitrogen,  increased  inversely  as  the  amount  of  sun- 
shine the  plants  received  we  would  expect  the  percentage  of 
protein  to  be  highest  in  the  plants  receiving  only  one  quarter  of 
the  total  sunshine  and  to  decrease  gradually  to  full  sunshine. 
This,  however,  does  not  seem  to  be  the  case.  Taking  the 
average  of  the  six  samples  grown  at  the  Tulare  station 
in  1908,  we  find  just  the  opposite  result.  The  protein  content 
increases  with  the  sunshine  quite  uniformly.  The  wheat  from  the 
plants  receiving  one-half  sunshine  contain  .47  per  cent  more 
protein  than  the  wheat  from  those  receiving  one-quarter  sunshine. 
The  wheat  receiving  full  sunshine  contained  .91  per  cent  more 
than  those  receiving  only  one-half  sunshine.  Unfortunately  the 
samples  grown  under  three-quarters  sunshine  were  lost,  so  that 
we  are  unable  to  say  whether  or  not  the  protein  content  here 
would  have  been  above  or  below  that  of  full  sunshine. 

The  analj^sis  of  the  individual  samples  in  this  set  show 
that  only  one  departs  markedly  from  the  average.  In  the 
variety  no.  879  we  find  that  the  wheat  grown  under  one- 
half  sunshine  had  a  higher  protein  content  by  .29  per  cent  than 
did  that  grown  under  full  sunshine,  while  that  grown  under 
one-quarter  sunshine  is  the  lowest  of  the  three  by  .17  per  cent. 
In  sample  no.  899  we  find  that  one-quarter  and  one-half  sunshine 
gave  nearly  the  same  results,  with  a  slight  difference  in  favor 
of  the  former. 

The  general  trend  of  the  results  from  1909  experiments  is 
much  the  same.  The  average  of  the  samples  grown  under  one- 
quarter  and  one-half  sunshine  are  about  equal,  while  the  protein 
content  of  those  grown  under  three  quarters  and  full  sunshine 
are  much  higher.  If  we  exclude  sample  no.  10-46G  grown  under 
one-half  sunshine,  which  is  unusually  low,  the  average  for 
one-half  sunshine  is  12.25  per  cent.  This  gives,  then,  quite 
a  gradual  increase  in  protein  content  from  one  quarter  to  three 
quarters  sunshine,  while  the  full  sunshine  samples  average  .5 
per  cent  lower  than  those  receiving  only  three-quarters  sunshine. 

The  results  from  the  experiments  at  Davis  are  slightly  more 
erratic,  but  the  general  trend  is  the  same  as  that  obtained  at 
Tulare  during  the  season  of  1909.  The  samples  under  one  third 
simsliine  average  .24  j)er  cent  more  protein  than  did  those  under 


1913]     Shaiv :  Studies  on  Influences  Afecting  Protein  Content  of  Wheat      109 

one-half  sunshine.  The  samples  under  two-thirds  sunshine  aver- 
aged .52  per  cent  more  protein  than  those  under  one-half  sunshine, 
and  .28  per  cent  more  than  those  under  one-third  sunshine. 
There  was,  then,  a  drop  in  the  full  sunshine  sample  to  about  the 
same  protein  content  as  those  receiving  one-half  sunshine. 

The  results  for  1909  at  Davis  came  in  the  same  order  as  those 
of  1908.  The  samples  under  one-third  sunshine  average  higher 
in  protein  than  those  under  t)ne-half  sunshine  by  .23  per  cent. 
The  samples  under  tw^o-thircls  sunshine  averaged  higher  than 
those  under  one-half  by  .6  per  cent,  and  than  those  under  one- 
third  sunshine  by  .37  per  cent,  w^hile  the  average  of  the  samples 
grown  under  full  sunshine  is  nearly  the  same  as  of  those  grown 
under  one-half  sunshine. 

These  results  certainly  show  that  the  protein  content  does  not 
vary  inversely  with  the  amount  of  sunshine  which  the  plants 
receive.  On  the  other  hand,  the  experiments  at  Davis  and  those 
of  the  year  1909  at  Tulare  tend  to  show  that  there  is  a  happy 
medium  under  which  the  maximum  amount  of  protein  is  stored. 
This  optimum  condition  seems  to  be  at  a  point  somewhat  below 
the  normal  sunshine.  If,  however,  the  amount  of  sunshine  falls 
below  that  medium  then  again  there  is  a  decrease  in  the  amount 
of  protein  stored.  Just  why  this  should  be  the  case  is  still  a 
matter  of  conjecture.  It  is  quite  probable,  however,  that  it  is 
due  to  a  disturbed  condition  of  the  physiological  functions  within 
the  plant  brought  about  by  the  abnormally  low  sunshine.  This 
fact  can  only  be  determined  by  a  closer  study  of  the  formation 
and  transformation  of  the  various  compounds  in  the  plant. 

The  gliadin  content  of  the  samples  in  this  experiment  seems 
to  bear  even  less  relation  to  the  sunshine  than  did  the  total 
protein.  In  fact,  there  was  no  regularity  whatever  in  the  results 
those  obtained  at  Tulare  being  just  opposite  from  those  obtained 
at  Davis. 

A  comparison  of  the  two  years  at  the  two  stations  shows  that 
there  was  a  marked  difference  in  the  protein  content  of  the  samples 
in  different  seasons.  At  Davis  we  find  that  the  samples  averaged 
a  higher  percentage  of  gluten  in  1909  than  in  1908  by  over  2  per 
cent. 


]  1 0         University  of  California  Pudlicatio7is  in  Agricultural  Sciences    [Vol.  1 

Comparing  the  weather  reports  at  Davis  for  the  months  of 
April,  May,  and  June,  we  find  that  this  period  during  1909  was 
much  drier  and  somewhat  warmer  than  the  corresponding  months 
the  year  before.  As  this  is  the  time  when  the  greatest  develop- 
ment of  the  kernel  takes  place,  it  is  not  at  all  unlikely  that  these 
climatic  differences  may  account  to  some  extent  for  the  dif- 
ferences in  composition  of  the  grain.  The  moister  and  cooler 
coi^dition  of  1908  may  have  prolonged  the  developing  and  rip- 
ening period,  thus  favoring  the  storing  of  a  larger  amount  of 
starch.  Other  seasonal  differences  not  recorded  have  doubtless 
also  contributed  their  share. 

The  difference  in  protein  content  for  the  two  seasons  at  Tulare 
is  just  as  marked  as  at  Davis,  but  the  order  is  reversed.  No 
weather  reports  from  the  immediate  vicinity  were  available,  so 
we  can  make  no  comparison  with  the  Davis  conditions. 

Another  rather  striking  seasonal  difference  brought  out  by 
the  table  is  the  percentage  of  gliadin  to  total  protein.  The  years 
that  the  percentage  of  total  protein  is  high,  the  percentage  of 
gliadin  to  total  gluten  is  also  high.  In  fact,  the  difference  in  the 
gliadin  content  of  the  wheat  samples  for  the  two  years  is  pro- 
portionally greater  than  the  difference  in  total  protein.  This 
would  lead  us  to  believe  that  the  gliadin  is  affected  more  by  the 
season  than  the  other  protein  compounds.  We  find,  however, 
little  or  no  relation  between  the  gliadin  content  and  the  sunshine 
received. 

In  conclusion,  then,  it  is  safe  to  say  that  while  sunshine  does 
exert  some  influence  upon  the  composition  of  the  wheat  grain, 
there  are  other  climatic  factors  which  also  exert  very  marked 
influences  in  this  direction.  We  find  that  the  protein  does  not 
increase  inversely  as  the  sunshine,  but  that  there  is  an  optimum 
condition  under  which  the  greatest  development  of  protein  takes 
place.  This  optimum  of  sunshine  is  somewhat  less  than  normal 
in  the  valleys  of  this  state.  Other  things  being  equal,  too  little 
sunshine  lowers  the  protein  content  to  just  as  great  an  extent  as 
too  much  sunshine.  This  condition  is  probably  due  to  the  fact 
that  a  certain  amount  of  sunshine  is  necessary  in  order  that  the 
normal  x>h.ysiological  functions  of  the  plant  may  take  place. 
When  the  amount  of  sunshine  is  reduced  to  one  quarter  or  one 


1913]     Shaw :  Studies  on  Influences  A  f  eating  Protein  Content  of  Wheat      111 

half  of  the  normal  it  is  quite  likely  that  the  plants  do  not  receive 
enough  sunshine  to  allow  even  the  maximum  nitrogen  metabolism 
to  take  place. 

The  fact  that  there  is  a  greater  difference  in  the  percentage 
of  protein  for  different  seasons  than  there  is  in  the  same  season 
under  various  amounts  of  sunshine  certainly  tends  to  show  that 
there  are  factors  other  than  sunshine,  which  play  just  as  im- 
portant a  part  in  determining  the  composition  of  the  grain. 

Certain  experimenters  have  stated  that  the  exposure  of  grain 
to  the  action  of  strong  sunlight  after  it  had  become  ripe  had  a 
tendency  not  only  to  bleach  the  kernels,  forming  the  so-called 
' '  yellow  berry, ' '  but  also  to  lower  the  protein  content.  Attention 
was  called  to  this  by  Lyon  and  Keyser  based  upon  some  trials 
made  at  the  Nebraska  station.  On  account  of  the  fact  that  the 
sunshine  in  the  main  grain-growing  regions  of  California  is  very 
intense  during  the  ripening  and  harvesting  period,  and  that  the 
grain  frequently  stands  for  several  weeks  on  the  straw  in  the 
field,  an  attempt  was  made  in  1906  at  Yuba  City  to  ascertain 
if  this  effect  held  under  the  conditions  which  obtain  in  this  state. 
In  this  experiment  several  varieties  of  wheat  were  selected  in 
which  differences  in  the  physical  appearances  of  the  grain 
could  be  easily  followed  owing  to  the  color  of  the  typical  and 
changed  kernels.  The  percentage  of  typical  kernels  in  the  orig- 
inal was  determined,  using  5000  kernels  in  each  case  as  a  basis, 
and  these  were  seeded  in  plats  under  like  conditions  in  the  field. 
At  the  maturity  of  the  grain  three  fourths  of  the  grain  was  cut 
while  in  the  hard  dough  stage,  a  few  bundles  being  shocked  and 
left  in  the  field,  while  an  equal  number  were  protected  from  the 
direct  action  of  the  sunlight.  The  remainder  of  the  grain  was 
left  on  the  standing  straw  from  July  5  to  August  13,  when 
it  was  harvested  and  all  three  lots  were  threshed  at  the  same 
time.  The  experiment  was  repeated  again  in  1907.  A  determi- 
nation was  made  of  the  percentage  of  typical  kernels  in  each 
lot,  and  of  the  percent  of  protein.    The  results  follow^ : 


112 


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1913]     Shaw :  Studies  on  Influences  Affecting  Protein  Content  of  Wheat      113 


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114         University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 

The  above  results  are  not  as  consistent  as  could  be  desired 
in  answering  the  main  question  involved  in  this  experiment  and 
no  attempt  will  be  made  to  interpret  them  as  bearing  upon  this 
particular  question.  They  do,  however,  further  bear  out  the 
results  discussed  in  the  experiment  on  the  effect  of  allowing  the 
grain  to  stand  on  the  straw  in  the  field  after  reaching  the  hard 
dough  stage,  for  it  will  be  noted  that  in  four  out  of  six  cases 
there  was  a  larger  percent  of  typical  kernels  in  the  late  cut 
samples  than  in  those  early  cut,  and  in  one  other  case  (no.  879) 
the  percentage  was  essentially  the  same  in  the  late  cut  sample 
as  in  that  early  cut.  In  the  matter  of  total  protein,  the  late  cut 
grain  all  carried  a  higher  percentage  than  did  those  of  the  early 
cutting. 

The  average  results  from  this  standpoint  are  shown  below : 

Per  cent  Per  cent 

typical  total 

kernels  protein 

Early  cut  69.1  12.92 

Late  cut  72.1  13.52 

THE  EFFECT  OF  IRRIGATION  UPON  THE  PROTEIN  CONTENT  OF 

WHEAT 

The  idea  has  been  quite  current  among  observant  growers 
that  whenever  the  rains  extended  late  into  the  spring  the  quality 
of  the  grain  of  that  season  was  materially  reduced,  that  this  had 
much  to  do  with  the  wide  seasonal  differences  in  the  quality  of 
grain  in  California,  and  possibly  was  the  main  factor  in  causing 
such  differences.  This  idea  is  quite  in  harmony  with  what  has 
been  observed  in  other  experiments  as  to  the  effect  of  irrigation 
upon  the  quality  of  grain.  No  definite  data  being  at  hand  as 
referring  to  conditions  in  California,  in  1908-09  trials  were  made 
with  six  types  of  wheat  at  Davis  to  determine  the  effect  of  early 
and  late  application  of  water  to  growing  wheat  by  planting 
these  six  types  on  uniform  soil  in  rows  at  about  the  ordinary 
rate  of  seeding  on  three  different  plats.  The  plats  received  the 
following  treatment  so  far  as  water  was  concerned: 

Plat  A  received  irrigation. 

Piat  B  wa.s  irrigated  in  the  rows  once  just  after  the  grain 
was  out  of  the  boot. 


1913]     Shaw:  Studies  on  Influences  Afecting  Protein  Content  of  Wheat      115 

Plat  C  was  irrigated  in  the  same  manner  twice,  once  at  the 
same  period  in  the  plants'  growth  as  in  Plat  B  and  again  just 
after  the  grain  set. 

All  plats,  then,  received  the  rainfall  of  the  season  and  Plat  B 
in  addition  had  one  irrigation  and  Plat  C  had  two  irrigations, 
the  last  one  very  late  in  its  period  of  growth. 

The  wheats  used  in  the  experiment  and  their  original  com- 
positions were  as  follows: 

Table  Showing  the  Analysis  of  Original  Wheats  Used  in  Irrigation 
Trials  at  Davis,  Cal. 


No. 
730/06 

Per  cent 
typical 
kernels 

96.0 

Number 

kernels 

in  10 

grams 

276 

Per  cent 

total 
protein 

12.83 

Per  cent 
gliadin 

4.97 

Per  cent 
ash 

2.32 

726/06 

87.7 

245 

13.55 

4.42 

2.20 

870/07 



227 

15.92 

4.27 

2.02 

049/08 

73.9 

313 

12.50 

4.24 

1.99 

870/07 

90.1 

239 

12.17 

4.12 

2.27 

338a2 

100.0 

284 

11.28 

2.13 

The  grains  were  seeded  on  December  7.  They  all  came  up 
with  a  good  stand  on  December  28-31  and  were  harvested  in 
two  lots  on  June  24  and  30,  there  being  but  one  or  two  days 
difference  in  the  time  of  ripening  between  the  irrigated  and  the 
non-irrigated  plats. 

The  analysis  of  the  several  lots  grown  on  each  plat  is  shown 
in  the  following  table: 


116         University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 


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1913]     Shaw :  Studies  on  In-fluences  Affecting  Protein  Content  of  Wheat      117 

Examining  these  results,  it  will  be  seen  that  in  both  the  early 
and  late  cut  lots  Plat  A,  which  received  no  late  application  of 
water,  carried  the  highest  average  per  cent  of  protein,  and  that 
Plat  C,  which  had  two  water  applications,  carried  the  lowest. 
Averaging  the  early  and  late  cut  lots,  the  following  figures  hold : 

Number 


A 

Per  cent 
typical 
kernels 

94.1 

kernels 
in  10 
grams 

240 

Per  cent 
protein 

14.70 

Per  cent 
gliadin 

5.22 

Per  cent 
ash 

1.85 

B 

88.1 

235 

13.78 

4.81 

1.94 

C 

83.1 

238 

13.40 

4.51 

1.84 

This  shows  a  gradual  decrease  in  both  the  protein  and  gliadin 
content  as  the  moisture  was  increased,  is  in  entire  harmony  with 
results  cited  elsewhere,  and  seems  to  show  that  either  irrigation 
or  late  rains  tend  to  lower  the  gluten  content  of  wheat,  and  that 
this  climatic  factor  is  a  very  prominent,  if  not  the  most  important 
one  in  causing  seasonal  variation  in  the  grain. 

Comparing  the  early  cutting  with  the  late  cutting,  it  will 
be  seen  that  while  there  is  some  fluctuation  between  corresponding 
samples  in  the  two  cuttings,  the  averages  bear  out  the  experi- 
ments cited  in  the  earlier  pages  to  the  effect  that  no  deterioration 
occurs  from  such  late  cutting,  and  as  a  matter  of  fact  in  this 
series  of  trials  there  was  an  actual  increase  in  the  protein  content 
in  the  late  compared  with  the  early  cutting; 

THE  EFFECT  OF  EEDUCING  THE  ATMOSPHEKIC  TEMPERATURE 
AT  NIGHT  UPON  THE  PROTEIN  CONTENT  OF  WHEAT 

In  the  season  1907-08  an  attempt  was  made  to  reduce  the 
temperature,  at  different  stages  of  plant  growth,  on  certain  plats 
on  which  were  seeded  several  types  of  grain  and  to  compare  the 
protein  content  on  these  plats. 

The  general  plan  of  this  experiment  consisted  of  seeding 
several  types  of  wheat  in  rows  upon  adjacent  small  plats  of 
uniform  soil.  The  rows  were  seeded  north  and  south,  and  to 
prevent  the  plats  from  receiving  the  early  sun  and  to  assist  in 
holding  down  the  temperature  in  the  early  morning  across  the 
south  end  of  the  plats  cooled  during  the  first  period  of  growth, 
a  board  fence  was  erected  sufficiently  high  to  shade  during  the 


118         University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 

morning  hours  about  one  half  of  each  plat  throughout  the  period 
of  cooling.  From  the  time  the  grains  were  well  up  until  the 
spring  rains  ceased  a  layer  of  ice  was  spread  on  a  loose  frame 
and  placed  over  one  half  of  each  plat,  within  four  inches  of  the 
top  of  the  plants,  every  night  during  the  first  half  of  the  growing 
period  of  the  plants.  A  portion  of  the  north  half  of  each  plat 
was  left  uniced.  To  retain  more  effectively  the  cooled  air  during 
the  night  the  iced  portion  of  the  plats  was  entirely  covered  by 
a  piece  of  heavy  canvas. 

In  this  experiment  two  varieties  of  common  wheats  and  five 
strains  of  durum  wheats  were  used.  The  composition  of  each 
sample  harvested  is  shown  below: 


Table  14. — Showing  Effect  of  Reducing  Night  Temperature  Upon  the 
Composition  of  Wheat  Kernels 


Not  Cooled 

Lab. 
No. 

745B/08 

Name 
Kubanka  1440 

Per  cent 
typical 
kernels 

98.65 

Number 

kernels 

in  10 

grams 

217 

Per  cent 
protein 

15.04 

Per  cent 
gliadin 

5.10 

Per  cent 
ash 

1.84 

851/08 

Kubanka  2221 

99.8 

201 

15.19 

5.09 

1.92 

869/08 

Kubanka  2239 

99.6 

177 

15.77 

5.44 

1.87 

871/08 

Velvet  Don 

99.5 

205 

15.44 

5.31 

1.96 

872/08 

Gharnovka 

99.5 

207 

15.15 

5.09 

1.83 

920/08 

Red  Fife 

96.9 

361 

17.40 

5.82 

2.19 

962/08 

White  Fife 

100.0 

323 

16.49 

5.65 

1.97 

Avera 

ge 

99.1 

241 

15.78 

5.36 

1.93 

Cooled  First  Period 


Lab. 
No. 

745B/08 

Name 
Kubanka 

Per  cent 
typical 
kernels 

99.75 

Number 

kernels 

in  10 

grnms 

254 

Per  cent 
protein 

14.95 

Per  cent 
gliadin 

4.52 

Per  cent 
ash 

1.88 

851/08 

Kubanka  2221 

99.70 

233 

15.31 

4.96 

1.23 

869/08 

Kubanka  2239 

100.00 

213 

14.61 

5.27 

2.04 

871/08 

Velvet  Don 

99.94 

229 

15.87 

5.32 

1.86 

872/08 

Gharnovka 

100.0 

241 

15.60 

4.67 

1.90 

920/08 

Red  Fife 

100.00 

384 

17.76 

2.26 

962/08 

White  Fife 

100.00 

380 

18.24 

5.75 

2.41 

Average 

99.9 

276 

16.05 

5.08 

1.98 

1913]     Shaw :  Studies  on  Influences  Afecting  Protein  Content  of  Wheat      119 
Cooled  Second  Period 


Lab. 

No. 

Name 

Per  cent 
typical 
kernels 

Number 

kernels 

in  10 

grams 

Per  cent 
protein 

Per  cent 
gliadin 

Per  cent 
ash 

745B/08 

Kubanka  1440 

100.0 

217 

15.24 

5.39 

1.96 

851/08 

Kubanka  2221 

100.0 

215 

15.45 

4.96 

1.80 

869/08 

Kubanka  2239 

99.8 

203 

15.80 

5.43 

1.90 

871/08 

Velvet  D 

99.9 

246 

15.53 

4.25 

1.90 

872/08 

Gharnovka 

100.0 

218 

16.15 

4.89 

1.90 

920/08 

Eed  Fife 

97.5 

350 

17.80 

5.96 

2.31 

962/08 

White  Fife 

99.4 

339 

17.69 

6.29 

2.13 

Avera 

ere 

99.6 

255 

16.23 

5.31 

1.99 

Collecting  the  averages  for  comparison,  the  figures  are  as 
follows : 

Number 
Per  cent       kernels 

typical  in  10  Per  cent  Per  cent    Per  cent 

Treatment  kernels  grams  protein  gliadin  ash 

Not   cooled  99.1  241  15.78  5.36         1.93 

Cooled  First  Period  99.9  276  16.05  5.08         1.98 

Cooled  Second  Period  99.6  255         16.23  5.31         1.99 


While  the  figures  for  individual  analyses  are  slightly  erratic, 
yet  it  appears  that  the  general  effect  of  reducing  the  temperature 
in  each  period  tended  to  increase  the  total  protein,  and  that  the 
tendency  was  greatest  when  the  night  temperature  was  reduced 
in  the  second  period  of  growth,  for  in  the  case  of  cooling  during 
the  first  period  of  growth  five  out  of  the  seven  trials  showed  a 
higher  percentage  of  total  protein  than  those  grown  under  normal 
conditions,  and  the  average  total  protein  was  16.05  per  cent 
against  15.78  in  the  case  of  the  uncooled  lot. 

In  the  case  of  the  lot  cooled  during  the  second  period  of 
growth  (after  the  grain  was  in  the  boot)  all  seven  showed  an 
increased  protein  content  over  the  uncooled  lot,  the  respective 
averages  showing  17.23  against  15.79  per  cent. 

Further,  the  effect  of  the  cooling  seems  to  have  been  greater 
from  reducing  the  temperature  during  the  second  period  of 
growth  than  during  the  first,  for  five  of  the  cases  in  this  com- 
parison show  increased  protein  with  an  average  protein  content 
of  16.23  per  cent  against  16.05  per  cent,  and  a  gliadin  content 
of  5.31  per  cent  against  5.08  per  cent. 


120  University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 

RELATION  OF   INCREASING   THE  SEVERAL   AVAILABLE   PLANT 

FOODS  IN  THE  SOIL  TO  THE  PROTEIN  CONTENT  OF 

WHEATS   AT   THE   UNIVERSITY  FARM, 

DAVIS,  1908-12,  INCLUSIVE 

The  general  effect  upon  the  protein  content  of  wheat  from 
increasing  the  available  nitrogen  and  other  plant  food  elements 
in  the  soil  at  the  University  Farm  at  Davis  has  been  made  the 
subject  of  study  for  the  past  four  years  and  the  results  are 
discussed  below.  In  these  experiments  Little  Club  wheat  has 
been  used  each  year.  In  order  that  any  cumulative  effect  which 
might  accrue  from  the  nitrogen  and  the  other  plant  foods  used 
might  be  apparent,  the  seed  from  each  plat  was  seeded  back 
upon  the  same  plat  each  succeeding  season. 

The  original  seed  used  in  these  experiments  was  grown  in 
1906,  and  had  the  following  composition: 


As  In  dry 

analyzed  matter 

Moisture  '. 11.28  

Total  protein 12.12  13.66 

Gliadin  4.38  4.93 

Glutenin  6.38  7.19 

Non-gluten  proteicls  1.36  1.58 

Ash  1.62  1.82 

Kernels  in  10  grams  — .        226 

Bushel  weight 59  lbs. 


For  the  four  years  1908,  1909,  1910,  and  1912  the  average 
result  was  as  stated  in  the  table  below.  During  the  season  of 
1911  the  land  was  under  bare  summer  fallow  on  account  of  its 
foulness  with  wild  oats,  which  accounts  for  the  omission  of  that 
year. 

During  these  trials  the  one-twentieth  acre  plats  received  the 
indicated  quantities  of  fertilizer  each  season. 


1913]     Shaiv :  Studies  on  Influences  Affecting  Protein  Content  of  Wheat      121 


Table  15.  Showing  the  Effect  of  Various  Fertilizer  Ingredients  Upon 
THE  Protein  Content  of  Wheat  Kernels 

Per  cent 
total         Per  cent 

Fertilizer  applied  protein         gliadin          Ash 

1.  Nitrate  of  soda,  5  lbs.;  hydrate  of  lime,  1321/2 

lbs *10.36         3.000         1.75 

2.  Hydrate  of  soda,  5  lbs *11.08         3.280         1.73 

3.  Hydrate  of  lime,  1321/0  lbs.  ...: 11.41         4.037         1.93 

4.  Cheek,  no  fertilizer  10.81          3.869          1.98 

5.  Nitrate  of  soda,  5  lbs.;  sulphate  of  potash,  6 

lbs ^ 10.99         3.816         1.95 

6.  Nitrate  of  soda,  10  lbs 11.00         4.130         1.99 

7.  Check,  no  fertilizer  11.18         3.866         1.86 

8.  Nitrate  of  soda,  5  lbs.;  superphosphate,  20  lbs.  11.02         3.855         1.82 

9.  Nitrate  of  soda,  10  lbs.;  sulphate  of  potash,  6 

lbs 11.91         4.416         1.80 

10.  Nitrate  of  soda,   10  lbs.;    superphosphate,   30 

lbs 10.93         3.728         1.83 

11.  Check,  no  fertilizer  9.99         3.793         1.87 

12.  Superphosphate,  20  lbs 10.64         3.888         1.86 

13.  Sulphate  of  potash,  6  lbs 10.82         4.054         1.81 

14.  Check   10.77         3.978         1.79 

15.  Nitrate  of  soda,  5  lbs.;  sulphate  of  potash,  12 

lbs 10.94         3.806         1.82 

16.  Nitrate  of  soda,  5  lbs. ;  superphosphate,  30  lbs.  10.57         3.878         2.20 

17.  Nitrate  of  soda,   10  lbs.;    superphosphate,   30 

lbs.;  sulphate  of  potash,  12  lbs 10.76         3.870         1.90 

18.  Nitrate   of   soda,    5    lbs.;    superphosphate,    30 

lbs.;  sulphate  of  potash,  6  lbs 11.00         4.291         1.87 

19.  Nitrate   of   soda,    5    lbs.;    superphosphate,    30 

lbs.;  sulphate  of  potash,  12  lbs 11.35         4.703         1.90 

20.  Nitrate    of    soda,    5    lbs.;    superphosphate,    50 

lbs.;  sulphate  of  potash,  6  lbs 11.92         4.815         1.91 

21.  Check   10.58         4.312         1.82 

22.  Sulphate  of  potash,  6  lbs.;  superphosphate,  30 

lbs 11.06         4.617         1.81 

23.  Dry    blood,    7    lbs.;    superphosphate,    30    lbs.; 

sulphate  of  potash,  6  lbs 11.24         4.748         1.86 

24.  Legume,    1907-08;    nothing,    1908-09;    super- 

phosphate, sulphate  of  potash,  1909-10  12.27         5.720         1.87 

25.  Superphosphate,  30  lbs.,  sulphate  of  potash,  6 

lbs.,    1907-08;    legume,    1908-09;    nothing, 

1909-10    12.86         5.440         2.03 

*  Plats  1  and  2  were  discontinued  in  1910  on  account  of  building  and  plats  28  and 
29  substituted. 


122         University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 

Table  15 — (Continued) 

Per  cent 
total  Per  cent 

Fertilizer  applied  protein         gliadin  Ash 

26.  Sulphate  of  potash,  6  lbs.,  superphosphate,  30 

lbs.,    1908-09;    nothing,    1907-08;    legume, 

1909-10    12.51         5.287         1.71 

27.  Check,  no  fertilizer  11.23         4.692         1.93 

28.  Nitrate  of  soda,  5  lbs.;  hydrate  of  lime,  132yo 

lbs *11.91         5.265         1.86 

29.  Nitrate  of  soda,  5  lbs *11.89         5.065         2.14 

*  Plats  1  and  2  were  discontinued  in  1910  on  account  of  building  and  plats  28  and 
29   were   substituted. 

THE  EFFECT  OF  NITROGEN 

Collecting  the  results  as  to  the  effect  of  nitrogen  upon  the 
protein  content,  the  following  figures  hold: 


Plats  receiving  nitrogen 

Check  plats 

Plat                      Per  cent 
No.                        protein 

1                          10.36 

Plat 
No. 

3 

Per  cent 
protein 

11.41 

2                          11.08 

4 

10.81 

Av.  for  2  yrs.  10.67 

11.11 

Plat                      Per  cent 
No.                        protein 

9                          11.91 

Plat 
No. 

5 

Per  cent 
protein 

10.99 

6                     11.00 

7 

11.18 

5                     10.99 

18 

10.82 

10                     10.93 

12 

10.64 

17                     10.76 

19 

11.35 

10                     10.93 

12 

10.64 

23                      11.24 

22 

11.06 

Average             11.10 

10.95 

From  the  above  it  does  not  appear  that  increasing  the  avail- 
able nitrogen  content  of  the  soil  in  these  trials  has  had  any 
material  influence  in  increasing  the  nitrogen  in  the  grain,  nor 
has  there  been  any  cumulative  effect  shown  by  its  use.  This  is 
shown  by  the  individual  cases  as  well  as  in  the  general  averages. 
Considering  individual  cases,  Plat  5  and  Plat  6  may  be  compared, 
each  receiving  equal  amounts  of  potash,  while  the  former  received 
also  100  pounds  of  nitrate  of  soda  per  acre,  but  it  carried  only 
.17  per  cent  more  protein  than  the  plat  receiving  no  nitrate. 
Phit  6  received  an  application  of  200  pounds  of  nitrate  of  soda, 


1913]     Shaw:  Studies  on  Influe7ices  Affecting  Protein  Content  of  Wheat      123 

while  Plat  7  received  no  fertilizer,  and  yet  the  average  protein 
content  of  the  latter  shows  .18  per  cent  higher  than  the  former. 
Further,  in  the  case  where  a  complete  fertilizer  was  used,  as 
in  Plats  17  and  19,  the  former  receiving  double  the  quantity 
of  nitrate  of  soda,  the  latter  showed  .59  per  cent  protein  above 
the  former.  The  results  for  the  entire  period  show  that  increas- 
ing the  available  nitrogen  of  the  soil  had  no  general  influence 
toward  increasing  the  protein  content  of  the  wheats,  either  when 
used  alone  or  when  used  in  connection  with  the  other  plant  food 
elements. 

Comparing  the  effect  of  nitric  nitrogen  with  organic  nitrogen, 
as  dried  blood,  in  a  complete  fertilizer,  the  following  results 
stand  for  the  four  years : 

1908 

Plat  18— Nitric  nitrogen  9.24 

Plat  23— Organic  nitrogen   10.28 

This  set  seems  to  show  a  very  slight  increase  due  to  the  organic 
nitrogen,  but  it  is  so  slight  as  not  to  affect  in  any  material  way 
the  quality  of  the  grain,  and  the  variations  are  such  as  to  render 
it  more  than  probable  that  such  variations  as  do  occur  are  due 
to  other  causes,  possibly  moisture,  rather  than  to  the  nitrate 
present. 

THE  EFFECT  OF  INCREASING  THE  NITROGEN  CONTENT  OF  THE 

SOIL  IN  THE  PRESENCE  OF  LIBERAL   AMOUNTS 

OF  PHOSPHORIC  ACID 

In  these  trials,  nitrogen  in  the  form  of  nitrate  of  soda  was 
used  in  connection  with  an  application  of  phosphates,  as  shown 
in  the  tabulation: 


1909 

1910 

1912 

Average 

11.32 

9.99 

13.45 

11.00 

10.77 

10.85 

14.08 

11.49 

itrogen  and  phosphate  plats 

Check  plats 

Plat                      Per  cent 
No.                         protein 

10                      10.93 

Plat 
No. 

8 

Per  cent 
protein 

11.02 

10                      10.93 

12 

10.64 

8                     11.02 

12 

10.64 

18                     11.00 

22 

11.06 

20                     11.92 

22 

11.06 

Average             11.3  6 

10.88 

124  University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 

The  average  difference  shows  .28  per  cent,  an  amount  which 
is  too  small  to  be  a  matter  of  consideration  for  practical  im- 
provement in  the  quality  of  the  grain.  Further,  it  will  be  noted 
that  such  differences  as  do  occur  did  not  run  always  in  the  same 
direction. 

THE  EFFECT  OF  PHOSPHATES 

In  these  trials  the  superphosphate  used  was  from  treated 
rock  and  the  amounts  used  are  given  in  the  scheme  previously 
shown. 

Superphosphate  plats 

Plat  Per  cent 

No.  protein 

12  10.64 

8  11.02 

10  10.93 

16  10.57 

20  11.92 

19  11.35 

22  11.06 

Average  11.07 

In  general,  the  phosphates  as  here  shown  appear  to  have  had 
a  slight  tendency  toward  increasing  the  total  protein,  but  it  is 
very  slight  on  the  average.  In  five  cases  out  of  seven,  however, 
the  phosphate  plats  carried  somewhat  higher  protein  than  did 
the  check  plats.  A  more  extended  study,  however,  will  be  re- 
quired before  any  definite  conclusion  can  be  reached. 

THE  EFFECT  OF  POTASH 

In  these  trials  the  potash  was  universally  used  in  the  form 
of  sulfate  in  the  amounts  indicated  in  the  general  tabulation. 


Check  plats 

Plat 
No. 

11 

Per  cent 
protein 

9.99 

2 

11.08 

6 

11.00 

2 

11.08 

18 

11.00 

15 

10.94 

13 

10.82 

10.84 

Potash  plats 

Check  plats 

Plat 

Per  cent 

Plat 

Per  cent 

No. 

protein 

No. 

protein 

5 

10.99 

2 

11.08 

9 

11.91 

6 

11.00 

13 

]0.82 

14 

10.77 

15 

10.94 

5 

10.94 

19 

11.35 

18 

11.00 

Aver 

age             ]  ]  .20 

10.96 

1913]     Shaic :  Studies  on  Influences  Afecting  Protein  Content  of  Wheat      125 

There  is  shown  a  slightly  increased  protein  content  on  the 
potash  plats  over  the  corresponding  checks.  Three  out  of  the 
five  cases  show  a  higher  protein  content  on  the  potash  plats  than 
on  the  checks,  and  one  shows  the  same  percentage  on  both  plats. 
'The  difference  is  so  small  in  the  average  that  it  might  easily 
be  due  to  difference  in  the  moisture  content  of  the  soil,  but  data 
along  this  line  as  relating  to  these  plats  are  not  at  hand,  and 
therefore  it  would  be  unwise  to  attempt  to  draw  conclusions 
until  such  data  are  secured. 

THE  EFFECT  OF  GREEN  MANURE  CROPS 

Plats  23,  24,  and  25  have  had  legumes  grown  upon  them 
every  third  year  and  cereal  crops  the  other  year  of  the  rotative 
period,  with  superphosphates  and  potash  added  the  second  year 
after  the  legume.  Comparing  the  result  from  these  plats  for 
the  short  period  the  trials  have  been  under  way  with  the  average 
of  the  check  plats,  and  with  the  average  of  those  on  either  side, 
the  following  results  appear: 

Per  cent 
protein 
Plats  with  legumes  11.06 

Average  of  all  checks,  no  legume  11.02 

Average  of  two  nearest  checks,  no  legume  .—     10.90 

It  is  not  apparent  that  any  material  change  in  protein  content 
has  been  effected  by  the  treatment  employed. 

GENERAL  CONCLUSIONS 

The  results  presented  in  the  foregoing  pages  seem  to  warrant 
the  following  conclusions : 

First — There  are  important  seasonal,  varietal,  and  individual 
variations  in  wheat  plants  with  respect  to  protein  content. 

Second — The  principal  factor  causing  the  most  pronounced 
variation-  in  the  protein  content  of  wheats  is  climate,  particu- 
larly the  moisture  supply  in  the  later  growing  period  of  the  crop. 

Third — The  tendency  of  wheat  kernels  to  change  from  a 
glutenous  to  a  starchy  condition  is  not  a  constant  one,  but  is 
mainly  dependent  upon  the  individuality  of  the  plant  and  upon 
seasonal  influences,  particularly  moisture  supply  in  the  latter 
part  of  the  growing  period  of  the  plants. 


126  University  of  California  Publications  in  Agricultural  Sciences    [Vol.  1 

Fourth — In  wheats  100  per  cent  of  which  are  entirely  starchy 
there  may  be  a  reversion  to  an  entirely  glutenous  condition  in 
a  single  season,  or  the  reverse  may  occur,  dependent  upon  the 
seasonal  condition. 

Fifth — Allowing  the  grain  to  stand  on  the  straw  in  the  field 
until  fully  ripe  does  not  materially  affect  the  protein  content. 

Sixth — The  protein  content  of  wheat  is  affected  by  the  time 
of  seeding,  the  product  of  late  seeded  grain  having  a  higher 
percent  of  protein  than  that  of  early  seeded  grain. 

Seventh — The  protein  content  of  wheat  is  very  largely  influ- 
enced by  the  water  content  of  the  soil  in  the  later  period  Ox  its 
growth,  and  the  effect  of  either  irrigation  or  rainfall  during  this 
period  is  to  lower  its  protein  content. 

Eighth — The  percentage  of  sunshine  which  the  grain  receives 
during  its  period  of  growth  has  a  somewhat  direct  bearing  upon 
its  protein  content,  but  other  seasonal  conditions  are  more  im- 
portant. 

Ninth — Retarding  the  growth  through  cooling  the  atmosphere 
has  a  tendency  to  increase  the  protein  content. 

Tenth — The  quantity  of  available  nitrogen  in  the  soil  either 
alone  or  in  the  presence  of  other  available  plant  foods,  provided 
there  be  sufficient  to  supply  normal  growth,  appear  to  have  little, 
if  any,  influence  upon  the  protein  content. 

Eleventh — The  low  gluten  content  of  wheats  grown  in  Cali- 
fornia is  not  due  to  soil  exhaustion,  but  rather  to  the  following 
causes:  (1)  To  climatic  factors  which  allow  a  long  growing 
period;  (2)  to  relatively  early  seeding;  (3)  to  the  use  of  varieties 
inherently  low  in  gluten;  (4)  to  a  lack  of  selecting  highly  glu- 
tenous seed. 

Acknowledgements :  The  author  is  greatly  indebted  for  valu- 
able assistance  rendered  in  the  prosecution  of  various  parts  of 
the  routine  analytical  work  which  furnishes  the  basis  for  this 
paper  to  Miss  Rachel  Corr  and  Messrs.  J.  D.  Rose,  B.  A.  Madson, 
and  E.  H.  Walters,  and  also  to  Messrs.  A.  J.  Gaumnitz,  H.  Fi 
Blanchard  and  J.  T.  Bearss  for  the  careful  attention  to  the  field 
production  of  the  samples. 

Transmitted  January  2d,  1913. 


